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Scientific Papers
Natural History Museum
The University of Kansas
31 March 2005 N u mber 38: 1 -27
Systematics of the Bufo coccifer Complex (Anura: Bufonidae) of
Mesoamerica
By
Joseph R. Mendelson IIP, Beck^ L. Williams-, CnKisTorHEK A. Sheil^
AND Daniel G. Mulcahv*
Ubi
' Dcpaitiuoit of Hci-pctologi/, Zoo Atlanta, S(HI Cherokee Ave SE, Atlanta, GA 3U315 \'' •*^^-dc\TY
-Departiiieut of Integrative Biologi/. University of California, Berkeley, C A 94720-iUO . .;;Vt-R^
^Departine)it of Biology, jolni Carroll University, University Heights, OH 44118
^Department of Biology, Utah State University, Logan^lT 84322-5305
CONTENTS
ABSTRACT 2
RESUMEN 2
INTRODUCTION 2
Acknowledgments 5
MATERIALS AND METHODS 5
Morphological Data 5
Molecular Data 6
ACCOUNTS OF SPECIES 6
Biifo coccifer 6
Bufo cycladen 11
Bufo ibarrai 13
Bufo pisinnus 15
Bufo portcri 17
Bufo signifer 20
MORPHOMETRIC ANALYSES 22
MOLECULAR ANALYSES 23
LITERATURE CITED 24
APPENDIX 26
© Natural History Museum, Tlu' University of Kansas ElTlSt MfiVf Lf'TarV '^^^ ^'" "'''■*'"^*^2
Museum of Compara*ve Zoology
Harvard Unlversitv'
] ds>^-
Scientific Papers
Natural History Museum
The University of Kansas
31 Mcirch 2003 Number 38:1-27
Systematics of the Bufo coccifer Complex (Anura: Bufonidae) of
Mesoamerica
ubrary
Joseph R. Mendelson III', Bhck^ L. Williams-, Christoi'hek A. Sheil\
ANL^ Daniel G. Mulcahv*
- ^m
' Dcimiiiiieiit of Ucrpctolo^ii, Zoo Atlniitu, S(H) Cherokee Ave SE, Atlanta, GA j03]5 H '-'•^y^-rrY
-Department ofhite^rative Biology, University of California, Berkeley, CA 94720-3140 V. ''c-^^^
^Department of Biolo<;y, John Carroll Universiti/, University Hei;^hti, OH 4411S
^Department of Biology, Utah State Universiti/, Logan, UT S4322-5305
CONTENTS
ABSTRACT 2
RESUMEN 2
INTRODUCTION 2
Al K\'OVVLEDGMENTS 5
MATERIALS AND METHODS 5
Morphological Da la 5
Molfxular Data 6
ACCOUNTS OF SPECIES 6
Biifo coccifer 6
Bufo cycladen 11
Bufo ibarrai 13
Bufo pisiinius 15
Bufo porteri 17
Bufo signifcr 20
MORPHOMETRIC ANALYSES 22
MOLECULAR ANALYSES 23
LITERATURE CITED 24
APPENDIX 26
© Natiiivil History Museum, The University of Kjnsds EmS^ MtiVf LF'TarV '^^^ ^'^' ' "^'■*""''*^2
Museum of Compara*ye Zoo'o^
Harvard Univ6rs[t>'
2 Scientific Papers, Naturai, Hisiorv Mi si,iiM,Tiii' Uni\i ksnv ui- Kansas
ABSTRACT Many populations of toads occurrinj^ between west-central Mexico and Panama have been
referred to Biifo coccifcr Cope, KSdd. While the taxonomic status of these populatitms has been c]uestioned for
many decades, a tliori>iii;h ie\ it'w ot the B. coccifer complex never has been presented. Based on evidence from
external morphology and a partial molecular data-set, we conclude that this complex consists minimally of six
species. Herein, we recognize B. ivicifcr Cope, 1866, B. cychidcii Lynch and Smith, 1966, and B. ibanai Stuart,
1954, and describe three new species.
Key Words: Bufonidae; Bufo coccifcr, Biifo cycladcn, Biifo ilnirrai, Biifo /'/s/mhhs, Bufo porlcri, Bitfo signifer;
Mesoamerica; taxonom\'.
KESUMEN Muchas poblaciones de sapos que se encuentian cntre el oeste-central de Mexico y Panama
han sido referidas como Bufo coccifcr Cope 1866. Aunc]ue el estado taxoncimico de estas poblaciones ha sido
cuestionado por muchas decadas, nunca se ha presentado una revision completa del complejo B. coccifcr. En
base a c\ idcncia do mortologi'a external y una base de datos moleculares parcial, concluimos que este complejo
consiste de al menos seis especies. En este trabajo, reconocemos B. coccifcr Cope, 1866, B. cycladcn Lynch and
Smith, 19h6, y B. ilnirrai Stuart, 1934, y describimos tres especies nuevas.
Palabraf Clave: Bufonidae; Bufo coccifcr, Bufo cycladcn, Bufo ibnrrai, Bufo pisnmus, Bufo portcri, Bufo signifer;
Mesoamerica; taxonomfa.
INTRODUCTION
The taxon Bufo coccifcr Cope, 1866 currently is applied
to toads that are distributed allopatrically in five regions
of Mesoamerica (Pig. I): (1) the Tepalcatepec Valley,
Michoacan, Mexico; (2) the Pacific slope of the Sierra
Madre del Sur in Guerrero, Mexico; (3) the southern side
of the Isthmus of Tehuantepec, Oaxaca, Mexico; (4) nearly
the entire Pacific versant from northwestern Guatemala
to northwestern Costa Rica; and (5) western Panama.
Additional records exist from the Atlantic versants of
Honduras and Nicaragua. The distribution of these
populations approximately matches the distribution
of low elevation tropical dry forest (Rzedowski,
1994; Campbell, 1999) in these regions. However, the
Rica: Cope, 1866:130) and suggested that the holotype
likely originated from somewhere on the Meseta Central
of Costa Rica; this referral was supported by Savage (1974).
Porter (1963) provided a range map and general diagnosis
of B. coccifcr Porter (1965) discussed the distribution of
the species in more detail, observing that the populations
in Michoacan, Mexico, and Oaxaca, Mexico, evidently
are allopatric — with a hiatus of approximately 140 km
between records from Oaxaca, Mexico, and records from
southeastern Guatemala. Stuart (1954a:20) referred to "a
chain cif Bufo coccifcr-Wke toads" distributed through the
subhumid habitats of Central America, and specifically
noted undescribed species in central Guatemala, and
population on the Pacific slope of the Sierra Madre del Sur another in Guerrero and the Isthmus of Tehuantepec
is an exception to this generality; these toads are found at
moderate elevations (ca. 1000 m) in relatively wet pine-
oak forest. (See Campbell and Duellman, 2000, for habitat
description.) Other records from more mesic, upland
habitats include the mountains of central Honduras
(McCranie and Wilson, 2002). An additional specie.s, B.
ibarrai Stuart, frequently has been assigned to the Bufo
coccifcr Group (e.g.. Frost, 1985). The taxonomic status
and distribution of B. ibarrai was reviewed by Mendelson
(2001); this species occurs primarily in upland pine-oak
habitats in Guatemala and is reported here lor the first
time in an adjacent region of Honduras.
With reference to the broad distribution ot lUdo coccifcr,
it has been suggested many times that these wirious
populations likely are not conspi-citic (Stuart, 1954a,
1963; Duellman, 1960; Porter, 1963, 1965; Zweifel, 1965;
McDiarmid .\nL\ I oster, 1981; Mendelson, 2001; McCranie
and Wilson, 2002). Dunn and Stuart (1951) commented on
region of Mexico. Stuart (1954b) subsequently described
the populations in the upland pine-oak zone of Guatemala
as B. ibarrai. Lynch and Fugler (1965) reported B. ibarrai
from Honduras. Subsequently, the taxon has had a long
period of uncertain status, typically being confused with
B. coccifcr (e.g., comments by J. A. Campbell in Frost,
1985:41). Meyer and Wilson (1971) placed B. ibarrai in
the synonymy of B. coccifcr. However, Mendelson (2001)
recognized B. ibarrai as a ciistinct species, described the
tadpole, and prcnided a new diagnosis and range map
of records tiom Guatemala; also, he suggested that
the species may be endemic to Guatemala. Based on
examination of specimens from Honduras, McCranie
and Wilson (2002) suggested that B. ibarrai does not
occur in that country. Lynch and Smith (1966) referred all
populations in Mexico to the new taxon Bufo cycladcn and
cited differences in the ad\'ertisemenl call (tidi' Poitei;
1965) and ecology of the Mexican and Central American
thestatusof the type locality for B. coccifcr (" hrnha" Costa populations. I lo\ve\'er, the morphologv of this t
ixon was
Toads oi- iiii- Bcro coccifir Comi'LLX
K ilometers
100
Fij;, 1 , A m>ip of MfsoaniL'iiLM, sluuving thu genoriili/ed di^li ibutioiis of fo.ids ivtc-ned to Biifo locci/ci soiimi Kito, .ind of /x ibaiiai (as recognizud
hi'iein).
diagnosed poorly, with respect to B. ibnrrai (see Mendelson,
2001 ) cind other popiilatiotis referable to B. coccifcr (Porter,
1967). Porter (1%7) criticized Lynch and Smith (1966) for
not supporting their claitns of timrphological differences
atnong the populations atid also claimed that there are
no real ecological differences among the populations
considered; Porter proposed this ta\on to be a mniicii
diibiiuii. The taxonotnic cofifiision surroiuidifig B. c\/cliu1cii
is evident iti the fact that the taxon appears Ofi some recent
checklists (e.g., Frost, 1985), but not oti others (e.g., Flores-
Villela, 1993; Catnpbell, 1999).
Se\eral authors have described the male ad \ertisefnent
calls from various areas. Porter (1964, 1965) described the
ad\'ertisement calls of B. cociifcr from the Tehuantepec
region of Oaxaca, Mexico, in addition to those of toacHs
he referred to B. coaifcr recorded in Guerrero, Mexico,
and several populations in Central Atnerica (El Salvador,
Honduras, Nicaragua, and Costa Rica; Porter, 1965). The
various descriptions and comparisons of advertisement
calls presented by Porter (e.g., 1964, 1965) are difficult
to interpret because he provided no voucher nufiibers
for recorded specimens, and the detail of his locality
information is inconsistent. When tracking down specific
localitv data provided by K. R. Porter, we found it useful
to refer to recording station descriptions presented in
his original, unpublished dissertation (Porter, 1962:table
1). Zweifel (1965) first reported the presence of toads he
referred to B. coccifcr in Panama and demonstrated that the
advertisement call of this population differed from those
of both the Mexican (= Isthtnus of Tehuantepec region of
Oaxaca) and other Central Americati recordings that were
published by Porter (1964). McDiarmid and Foster (1981)
described additional advertisement calls recorded in Costa
Rica, and compared them to those reported from Mexico
and Central America (Porter, 1965) and from Panama
(Zwiefel, 1965); there are substantial differences in pulse
SciENTiiic Papi:rs. Natlrai. Hisiorv MusiiUM.Tiiii UNi\i;Rsir-t oi Kansas
labk' 1. Ciimporison ot i\ingi"s or moiins ot c\ills of the Hufo coccifcr ciimplex Ironi dlllcrcnt populations. Tlic identity ot the toads recorded
from Honduras (Porter, 1965) are unknown because the original paper provides neither reference to voucher specimens nor a detailed localilv-
description; thiTc ,ire at least three species in this group in ?londuras.
.\o. individuals
Duration
Frequency
Pulse Rate
Species
Location
recorded
(second)
(Hertz)
(per second)
Source
identity
Mexico: Guerrero
1(6 calls)
mean =5.7
mean =2800
mean =120
Porter, 1965
B. cycladcn
Mexico: Oaxaca
18
1.4-6.4
2800-3350
97-115
Porter, 1964, 1965
B. coccifcr
El Salvador
14
3.0-16.3
2200-2700
80-101
Porter, 1965
B. coccifcr
Guatemala
7
4.2-5.8
1650-1800
48-61
Porter, 1966
B. ilmrrai
Honduras
2
5.3-8.8
2200-2500
80-96
Porter, 1965
unknown
Nicaragua
4
6.2-10.9
2300-2400
91-100
Porter, 1965
B. coccifcr
Costa Rica
4
1.5-10.0
2300
90-95
Porter, 1965
B, coccifcr
C\>sta Rica
1
1.2-5.7
2231-2539
90-105
McDiarmid &
Foster, 1981
B. coccifcr
Panama
1
8.0-17.0
2100-2600
66
Zweifel, 1965
B. sigtiifer
rates and dominant frec|Liencies among these samples.
Porter (1966) described the ad\'ertisement call of B. ibiinui.
The data reported bv I^orter, Zweifel, and McDiarmid and
Foster are presented in Table 1 and summarized in Figure
2. We are unaware of any recordings of populations of 6.
cf. coccifcr from Guatemala or Michoacan, Mexico.
The calls of male toads from the Sierra Madre del Sur
of Guerrero and the Tehuantepec region of Oaxaca have
higher dominant frequencies than calls of all other samples.
The call of the toad in Guerrero also has a higher pulse rate
than all other reported calls. Note, however, that the data
from Guerrero are based on a recording of six separate
calls by the same individual. These data were used, in
part, by Lynch and Smith (1966) to justify recognition of all
Mexican populations (including populations in Michoacan,
Guerrero, and Oaxaca) as a species (B. cyclndcii) distinct
Dominant Frequency
Guerrero
Oaxaca
Central America
Panama
B. ibarrai
1600
2500
Hz
~1
3400
Pulse Rate
Guerrero
Oaxaca
Central America
Panama
B. ibarrai
40
I
80
pulse/sec
120
Fig. 2. Cirapliical representation ol Irequencies ami pulse rate's ot advertisement calls Ironi toads ul Ihe Biito coccifcr complex Ironi dilterent
regions of Mesoamerica; information shown here are a graphical representation of data presented m l.ible I IXila from "Cenlral America" here
includes range of values representing samples from El Salvador, Nicaragua, and Costa Rica.
Toads of thi-; Bufo cocciii-r Comi'i k.\
expressed or implied consensus of these authors is
that a thorough revision of the Biifo coccifcr complex is
long overdue, in tliis paper we use data from external
morphology and male advertisement calls (previously
published) to delimit species boundaries amting samples
referable to the widespread taxon B. coccifcr. We also present
a preliminary assessment of phvU>genetic relationships
among the species in the group, based on mtDNA data
from several of the species.
Acknowledgments
We are grateful to the following inclividuals for
providing assistance in the field, laboratory, and library: W.
Duellman, S. Cotte, K. Lips, M. Acevedo, E. Greenbaum,
J. R. McCranie, J. Malone, D. Laurencio, M. Sasa-Marin,
M. Ryan, M. Forstner, T. Reeder, J. Campbell, E. Smith,
C. Franklin, and R. CJutberlet. Adele Cutler provided
statistical acivice and assistance. Michelle Koo graciously
provided access and assistance with mapping software at
the California Academy of Sciences. We benefited greatly
from memories, observations, and notes drawn from the
considerable fieki experiences of J. Campbell, C. Myers,
j. Sa\'age, W. Duellman, anci R. Zweifel. Permission and
help with photographs was provided by M. Sasa-Marin,
R. Zweifel, and J. Simmons. Loans of critical specimens,
for extended periods of time, are gratefully acknowledged
from the curators and collection managers of the
following institvitions: AMNH, CAS, FMNH, TCWC,
UTA, TNHC, ANSP, UIMNH, USNM, KU, UMMZ,
MVZ, LACM, and LSUMZ. Some of the fieldwork for
this project was supported by funds from The National
Geographic Society and this research was conducted in
direct association with The Research Analysis Network
for Neotropical Amphibians (RANA; NSF-DEB 0130273).
Comments on the manuscript were provicied by the
students and faculty of the USU Herpetology Group, L.
frustration in delimiting species in this complex, which Trueb, J. Pramuk, and W, Duellman. Additional help and
is characterized bv advertisement calls that vary among favors were provided by J. Meik and G. Schneider,
populations and subtle morphological differences. An
MATERIAL AND METHODS
from B. coccifcr in Central America. A call described by
Zweifel (1465) from Panama differs dramatically from all
others b\' having a much lower pulse rate. The relationship
between the calls from Oaxaca and Central America is
interesting in that they have distinctly different ranges
of dominant frequencies and somewhat different (but
o\'erlapping) ranges of pulse rates. However, we note
that the Central America data do not include recordings
from Guatemala — the closest geographic region southeast
of the Oaxacan population. Recordings from Guatemala
may tiiminish the apparent distinctit)n between the
advertisement calls from the Tehuantepec region of Oaxaca
and Central America. Porter (1966) described the call of B.
ibarrni from Guatemala; this species differs from all other
populations in the B. coccifcr complex by having a very low
frequency and pulse rate (Table 1).
In their description of Bufo ci/chhicii, Lynch and Smith
(1966) placed great diagnostic importance on variation in
calls between Mexican and Central American populations.
We note that their summary of calls from Mexico was a
composite of data from both Guerrero anti the Tehuantepec
region of Oaxaca (data published by Porter, 1964, 1965; Table
1 ). VVe have concluded that these toads from Guerrero are
not conspecific with those from Oaxaca (discussed below).
Gergus et al. (1997) described call variation among toads of
the B. Diicroscnpliufi complex; they reported wide variation
in some parameters and an overall pattern of apparent
plesiomorphic similarity among allopatric species. There is
no reason to expect drastic differences in mate-recognition
systems among allopatric members of a complex of closely
related anurans with similar natural histories (Gergus et
al., 1997). Nevertheless, there are apparent differences in
the advertisement calls of toads of the B. coccifcr complex
that suggest diagnostic differences in the mate-recognition
systems of these allopatric populations.
All previous workers (above citations) have expressed
General terminology and measurements are those of
Mendelson (1997). Adult males were identified by the
presence of vocal slits and nuptial excrescences; large
individuals lacking these characters were presumed to be
adult females. If sex could not be determined externallv,
it was verified by direct observation of the gonads.
Foot-webbing formulae follow the system of Savage
and Heyer (1967), as modified by Myers and Duellman
(1982) and Savage and Heyer (1997). The general format
of the descriptions and diagnoses is slightly modified
from that of Mendelson (2001). Museum codes are those
proposed by Leviton et al. (1985). We follow Tyler et al.
(2001) in OLu- usage of the term "parotoid gland." Data
from advertisement calls were taken from the literatiuv
(referenced below).
MORPHOMI^TRIC DaIA
The following measurements were taken from adult
specimens: snout-vent length (SVL); head length (HL);
head width (HW); tibia length (TIB); foot length (FL); width
of tympanum (TYM); length of parotoid gland (PARE);
maximum width of parotoid gland (PARW); and length of
supratympanic crest (SPTYM). These variables represent
repeatable morphological landmarks and were measured
6
Scientific Papers, Natural History Museum. The University oi- Kansas
with digital calipers and iDiinded tt) the nearest 0.1 mm.
Because of the paucity of large series of females from many
localities, all morphometric analyses are based only on
adult males. Many of the data for Biifo ibnrnii are the same
presented by Mendelson (2001), but supplemented with
additional material frc)m both Guatemala and Honduras.
We conducteci a Principal Components Analysis
(PCA) using the covariance matrix on log-transformed
morphometric measurements from 324 adult male toads;
this data-set included specimens representing all species
and geographic regions of the distribution of the Bufo coccifcr
complex. We also performed a PCA on the residuals of the
latter seven variables derived from a regression analysis
(using SVL as the independent variable). This type of PCA
removes the effect of size, and displays general variation
in shape and proportion among the specimens (Good and
Wake, 1992). We performed a Linear Discriminant Function
Analysis (LDA) on the same data-set, and used residuals
of variables regressed on SVL and log-transformed SVL.
In this analysis, a priori groupings corresponded to the
species recognized in this paper (Accounts of Species,
below). Statistical analyses were performecl using SAS
and Minitab software.
Molecular Data
We conducted molecular analyses on ail available
samples, which inckided five specimens of Bufo coccifer (4
from Central America, 1 from Guerrero, Mexico) and two
specimens of B. ibnrrai. Additionally, one specimen of each
of the following species was included as outgroup taxa:
B. conifcrus, B. valliceps, and B. mnrijius. The selection of
these outgroup taxa was based on preliminary analyses of
a data set containing approximately 25 species of Central
American bufonids (Mendelson and Mulcahy, in prep.).
Sections of tiie mitochondrial genes cytochrome-b (cyt-b)
and 16S were used in the molecular analyses. Isolation and
PCK amplification of the mitochondrial DNA (mtDNA)
genes were performed exactly as described in Mulcahy
and Mendelson (2000), which includes primer information
and amplification prt)files. Products from the PCR were
amplified and sec]uenced in both directions using BigDye™
Terminator Cycle Set]uencing Ready Reaction Kit (Applied
Biosystems Part No. 4303152); we used the same PCR
primers and standard sec]uence-reaction profile on a Perkin
Elmer GeneAmp 2400 cycle sequencer. Cleaned sequences
were then run on an ABI 377 automated sequencer by
DGM at the Biology Department at Utah State University.
Sequence comparisons and alignment were conducted
with Sequencher 3.1.
Phylogenetic analyses of the mtDNA sequences
were conducted using PAUP* 4.0b8a (Swofford, 2002).
A partition-homogeneity test of 100 replicates was
implemented in PAUP (using default parsimony settings,
with the exception of addition-sequence being random
with 100 replicates) between cyt-b and 16S gene regions to
determine whether or not the two genes yielded conflicting
results. Maximum-parsimony analyses (MP) were
performed on each gene separately and both combined.
The program Modeltest (Posada and Crandall, 1998) was
used to evaluate the best Maximum-likelihood model, using
the Hierarchical Likeliht)od Ratio Tests (hLRTs) criterion.
A Maximum-likelihood (ML) analysis was then conducted
using the model and settings based on the hLRTs results.
Because of the limited number of taxa in the phylogenetic
analyses, an exhaustive search was possible, and used in
the MP analysis, while the ML analysis required a heuristic
search algorithm. Gaps were treated as "missing data,"
with characters-state optimization set at ACCTRAN.
Branch support was assessed bv nonparametric bootstrap
analyses using 1000 replicates of full heuristic searches,
with 100 random additions at each replicate, under MP and
100 replicates of full heuristic searches, with 10 random
additions under the ML criteria. Decay indices (Bremer,
1994) were also measured under the parsimony analysis
using the program TreeRot (Sorenson, 1996).
ACCOUNTS OF SPECIES
We used data from adyertisement calls, nu)rphometry,
external morphology, and DNA sequences to examine
variation among samples of the Bufo coccifcr complex.
These data are consistent with the "chain of Bufo
coccifer-Vikc toads" distributed through the subhinnid
habitats of Central America that Stuart ( 1954a) envisioned.
Evaluation of our data with respect to the Evolutionary
Species Concept (sensu Wiley, 1978; Frost and Hillis,
1990) supports recognition of six species in this complex.
Our proposed taxonomy reflects the sentiments of the
many authors (e.g., Stuart, 1954a, 1963; Duellman, 1960;
Porter, 1963, 1965; Zweifel, 1965; McDiarmid and Foster,
1981; Mendelson, 2001; McCranie and Wilson, 2002) who
have dealt with these toads during the last five decades.
We provide species accounts and diagnoses for B. coccifer,
B. ci/cladcn, and B. ibnrnii, and describe three new species
from this complex; photographs of these species in life are
presented in Figure 3.
Bufo coccifcr Cope
Figs. 3-5
Bufo coccifcr Cope, 1866:00. Hiil.it\pc: USNM 6490. Type locality:
"Arriba" Costa Rica.
lUifo coccifcr — Dunn and Emlen, 1932; Kellogg, 19,12; Hartweg and
(.liner, 1940; Dunn and Stuart, 1951; Smith and Taylor, 1948 [in part
Mertens, 19.S2; Taylor, 1932; Stuart, 1954b; Rand, 1957; Duellman, 1960
I'orter, 1963 |in part); Stuart, 1963; Porter, 1964; Porter, 1965 (in part]
Toads of thh Bufo coccifer Complex
Fig. 3. Species of the Bi{fo coccifer complex in life, clockwise from upper left: B. coccifer from Santa Rosa, Costa Rica (adult male: photograph
hy Andrea Bernecker); B. cyclndcii from Guerrero, Mexico (adult male, UTA-JRM 4607; photograph by ]. R. Mendelson); B. ibarrni from Baja Verapaz,
Guatemala (female, KU 186304; photograph by J. A. Campbell); B. pifimiiif from Michoacan, Mexico (adult male, from UMMZ series; photograph
bv W. E. Duellman); B. portcri from Francisco Morazan, Honduras (subadult female, KU 103220; photograph bv W. E. Duellman); B. si\;uifcr (adult
female, AMNH 69625; photograph by R. G. Zweifel).
Sfii:Nrii i( P,\i'i-.ks. Nvn RAi, lIisroK^ Mii.si-;um,Thk Univhrsii'i oi- Kansas
Lvnch and Fuglor, 1965; Porler and Porter, 1967; Meyer and Wilson, 1971
|in part); Villa, 1972; Savage, 1974; McDiarmid and Foster, 1981; Villa,
1983; Frost, 1985 |in part]; Savage and Villa, 1986; Villa et al., 1988; |in
parti; Campbell and Vannini, 1989 [in part]; Flores-Villela, 1993 |in part];
Campbell, 1999 |in part]; Kiihler, 1999; Mendelson, 2001; Frost, 21)03 |in
parti; McCranie and Wilson, 2002 |in part]; Savage, 2002.
Biifo cifcliuleii — Lynch and Smith, 1966 [in part, for reference to
specimens from Tehuantepec region of Oaxaca, Mexico]; Frost, 1985 ]in
part]; Frost, 2003 ]in part],
Biifo iharrai — Lynch and I'ligler, 1965 [in part].
Bufo vatlicepf micwli^ — Werner, 1896 (synonyiiu' b\' Mendelson,
2001).
Diagnosis. — A medium to large species of Bnfo
{males to 62 mm SVL; females to 82 mm SVL) having
the following combination of characters: (1) tympanum
e\'iclent externally, about 35-45' V' diameter of orbit in males
and females; (2) cantlial, supraorbital, supratvmpanic,
postorbital, preorbital, pret\'mpanic, parietal, and
siipralabial crests present; (3) cranial crests well dexeloped,
i(.)biist, except parietal mav be thin, low, or absent in some
specimens; (4) tibia short, about 35% SVL; (5) feet short,
about 35% SVL; (6) dorsal tubercles small to medium
sized, elevated, rounded, scattered relatively sparsely
on middorsal, dorsolateral, and lateral regions of bod\'
beci)ming more denseh' arranged and distinctly spinose
laterally in specimens from most regions; (7) yentral
tubercles granular, smooth, or with spinose apices; (8)
lateral descending row of enlarged tubercles absent; (9) skin
texture not sexually dinn)rphic; (U)) \ocal slit unilateral in
male; (11) in. iiitcrln/oidciii^ poorly diflerentiated from ;;;.
iiitciiiiaiidibiilnris, but differentiated posteriorh' forming a
large, unilobed vocal sac with heavy black pigmentation;
(12) snout rounded in lateral profile, pointed in dorsal
aspect; (13) parotoid glands round to oxoid and large, about
Fig. 4. Dorsal and ventral aspiTts ol representalise adult spi-ciniens of Hufo coctifcr from Ketalhiileu, C.iiatcniala (male, letl: UT.'\ .A-2,5821, SVL
= 57.0 mm; female, right: UTA A-29025, SVL = 75.0 mm) and B. ci/chulcn Irom t.iierrero, Mexico (male, lelt: L'MM/ 1 1 5357 ] WHO 92751, SVL = 53.8
mm; female, right: UMMZ 119270 ]WKD 1.3425], SVL = 61.4 mm).
ToAUs oi- iiii-; Brio coccii i:k Comi'lilX
Fig. 5. L.itei mI .ispccts ot tlif hiMd'i i.)l adult nulf >pf cimens of Biifo
coccifer (upper: KU hS4(l(l) .ind B. cycln'icu (lower: KU 97434). Scale bars
= 1 cm.
1.0-1.5 times size of eyelid; (14) skin between cranial crests
on top of head with few to many scattered, low, rounded
tubercles; (15) ventral coloration whitish cream, sometimes
with some degree of diffuse dark pigmentation, stunetimes
in the form of diffuse punctuations.
Specimens referable to Biifo coccifer vary considerably
in size, shape, coloration, skin texture, and parameters of
the advertisement call across the range of the species. Biifo
coccifer differs from B. ci/cladeii by: being larger (males of B.
cyclndcii to 54 mm SVL, females to 62 mm); having dorsal
tubercles that are relatively small and scattered about the
dorsal surfaces, rounded on dorsum, becoming spinose
laterally in most specimens (tubercles large, elevated,
and densely arranged in B. cycliiden, rounded dorsally,
becoming very spinose laterally); ventral tubercles that
are smoothly granular or with small, spinose apices
(ventral tubercles in /-!. c\/clih1eii with distinct, large
spinose apices); relati\el\' well de\eloped parietal crests
that rarely are absent (parietal crest weakly de\'eloped in
B. cyclndeu, sometimes absent); and an advertisement call
with a lower pulse rate (Table 1; Fig. 2). Biifo coccifer can be
distinguished from B. pisiumts by: being larger (males of
B. yisiuiiiifi to 51 mm SVL, females to 62 mm); and having
skin tubercles that are overall larger (all tubercles minute
in B. pi:^iuiiiis). Biifo coccifer may be distinguished from B.
signifer by having: no, iir few, relatively indistinct dark
brown markings on the venter (that of B. i^igiiifcr boldly
marked with a marbled pattern); relatively thicker and
higher cranial crests (all crests relatively low and thin
in B. signifer); and an advertisement call with a higher
frec^uency and pulse rate (Table 1; Fig. 2). Biifo coccifer
differs from /->. ihnrnii by: being smaller (males of B.
ibarrai to S2 mm SVL, females to 94 mm); having rounded
to subovoid parotoid glands about 1.0-1.5 times size of
eyelid (parotoid glands distinctly ovoid, about 1.5-2.0
times size of eyelid in B. ibarrni); and males usually with
dorsal tubercles becoming spincise laterally and females
with rounded tubercles middorsally (dorsal tubercles of
males of B. ibarrai rounded laterally and females with
spinose middorsal tubercles). Bufo coccifer differs from
B. porteri by having: sharply spinose lateral tubercles in
males (rounded in males of 6. porteri); and a relatively
thin supratympanic crest (large, bulbous in B. porteri).
Morphometric variation is summarized in Table 2.
Distribution and Ecology. — Bufo coccifer occurs along
the Pacific versant of Mesoamerica, from the Guatemala-
Mexico border to the Guanacaste region of Costa Rica
(Fig. 6). Additional records exist from interior valleys on
the Atlantic versant of Honduras and from the Atlantic
coastal regions of Honduras and Nicaragua. McCranie and
Wilson (2002) erroneously reported this species from the
Atlantic versant of Guatemala; there are no records from
this region. An apparently isolated population occurs in
the southwestern region of the Isthmus of Tehuantepec,
Oaxaca, Mexico. This species ranges from sea level to 1435 m
(Savage, 2002; record from Cartago, Costa Rica) and occurs
in a variety of tropical dry forests and savanna habitats.
Duellman (I960) provided photographs of the habitat
and some natural history observations from the Isthmus
of Tehuantepec. McDiarmid and Foster (19cSI) described
the reproductive biology of a population in northwestern
Costa Rica. The ecological notes provided by McCranie
and Wilson (2002) for Hondiu'an populations represent
mixed observations pertaining to B. coccifer, B. ibarrni, and
B. porteri. Savage (2002) provided a concise summary of
the ecology of this species, based on observations from
Costa Rica.
Tadpole. — McDiarmid and Foster (1981) and Savage
(2002) described tadpoles from Costa Rica and provided
illustrations.
10
Scientific Papi-ks. Nau kai History MLisi-.uM.THi-. Univi-ksha oi Kansas
Table 2. Morphometric variation among males and femak's from of the Biifo ccccifcr complex; mean + SD is presented above the range (in mm).
Data for B. ibarrai taken, in part, from Mendelson (21)01 ).
S. cocciff)
B. cydadcn
B. ibarrai
/' yi'-nniu-^
/ ■'
:: <.-:lfcr
Males
u = 130
n ^ 311
u ^ 79
n = 2l
It =43
» = 21
SVL
51.4 ±4.8
51.1 ±2.2
62.2 ±9.1
45.9 ± 3.2
49.6 ± 4.6
51.2 ±64
41.0-64.4
47.4-54.2
42.5-82.4
38.5-51.4
40.2-58.7
42.0-64.1
Tibia length
18.3 ±1.9
18.4 ±0.9
24.1 ± 3.6
15.9 ±1.4
18,4 ±1.8
17.9 ± 2.3
13.3-22.5
16.6-20.1
15.7-31.8
13.0-18.0
15.2-21.7
15.0-23.4
Foot length
18.1 ± 1.9
18.3 ±0.7
24.6 ± 3.8
16.9 ±1.3
18.8 ±1.8
18.2 ± 2.3
13.1-23.9
16.9-19.8
16.3-33.4
14.8-19.2
15.7-22.7
13.7-23.1
Head length
17.8 ± 1.8
18.0 ±0.9
20.8 ± 2.8
15.1 ± 1.0
16,7 ±1.4
174 ±2.0
13.6-21.6
16.7-20.9
15.3-26.5
12.9-17.1
14.0-20.0
14.7-22.2
Head width
20.5 ± 2.4
20.1 ±0.9
23.9 + 3.1
17.3+1.4
18.9 ±1.6
19.8 ± 24
15.9-27.8
17.6-21.8
16.2-30.8
15.1-20.2
15.7-23.0
16.6-24.9
Tympanum
2.3 ± 0.5
2.7 ± 0.3
3.7 ± 0.7
2.3 ± 0.3
3.0 ± 0.4
3.1 ± 0.4
1.3-3.7
2.1-3.3
2.7-5.3
1.8-2.6
2.2-3.5
2.5-3.8
Supratympanic
3.2 + 0.5
2.7 ± 0.3
3.5 ± 0.5
2.6 ± 0.3
3.1 ±0.3
3.0 ± 04
crest
2.0-4.4
2.2-3.3
2.3-5.2
2.2-3.2
2.3-3.7
2.1-3.8
Parotoid length
6.3 ±1.0
6.9 + 0.5
7.8+1.5
5.4 ± 0.6
6.3 ± 0.9
6.3 ±1.2
4.3-8.7
6.0-4.8
3.2-11.0
4.1-6.5
5.0-9.0
2.8-8.8
Pan>toid width
6.6 ±1.3
6.1 ±0.6
5.7 ± 0.9
5.3 ±0.5
5.1 ±0.7
6.0 ± 0.8
3.7-9.5
5.0-7.2
3.5-7.4
4.5-6.2
3.8-6.7
4.6-7.7
Females
It = 25
n =6
n = 61
» = 5
// = 14
11 = 11
SVL
67.7 ± 7.6
59.8 + 2.8
78.4 ± 7.6
59.1 ± 3.3
64.5 ± 5.8
67.2 ± 8.3
58.1-82.4
54.6-62.9
60.8-94.4
55.4-62.2
56.4-73.0
54.5-77.0
Tibia length
22.2 ± 2.4
20.0 ±1.6
28.7 ± 3.0
18.7 ±1.1
77 5 + 2.2
23.0 ± 3.4
18.8-28.1
18.3-22.3
20.2-32.8
18.0-20.6
18.7-27.2
17.7-26.5
Fool length
21.9 ±2.2
20.4 ±1,0
28.7 ± 3.7
20.3 ± 1.3
22.5 ±1.9
23.1 ±3.7
17.9-27.2
19.4-22.1
19.6-34.1
18.8-21.7
19.6-26.5
17.0-26.8
Head length
?? ? ± 2.2
20.7 ±7.6
25.2 ±2.1
18.8 ±0.9
20.9 ± 2.0
22.6 ± 2.5
19.2-26.8
18.9-22.8
19.8-28.5
17.6-19.7
18.0-25.9
18.4-25.9
Head length
25.5 ±2.5
23.0 ±1.5
29.5 ± 2.8
21.9 ±1.2
23.9 ±1.9
26.1 ±34
22.5-31.5
21.1-24.4
21.5-33.3
20.6-23.7
21.5-28.3
20.9-29.4
Tympanum
3.1 ±0.6
2.8 ± 0.5
4.1 ±0.5
3.2 ±0.1
3.5 ± 0.5
3.6 ± 0.4
2.1-4.2
2.2-3.5
3.0-5.4
3.1-3.4
2.6-4.5
3.1-4.3
Supratvnipanic
4.3 ± 0.8
3.1+0.2
4.5 ± 0.7
3.4 + 0.3
3.9 ± 0.6
4.0 ± 0.7
crest
2.9-6.2
2.8-3.4
2.9-6.0
3.1-3.8
3.0-5.2
3.1-5.2
Parotoid length
7.4 + 1.3
7.6 + 0.8
8.4 ±1.5
6.8 ±1.0
8.3 ± 08
7.8 ±1.3
5.4-9.9
6.5-8.5
3.0-11.7
5.9-8.4
6.8-9.9
5.5-9.9
Parotoid width
7.4 + 1.1
6.9 ±0.8
6.3 ± 0.9
6.6+ 1.0
6.6 + 0.4
7.7 ±1.0
■i 7 1114
6.2-8.4
4.1-8.2
■^ (. ~ s
b 1 - 2
5.4-9.0
Remarks. — Ci)k>r photographs ot this species appear
in Leenders (2()01:pl. 7), Savage (20n2:pl. 77), and Villa
(1972:fig. 57). Blair's (1963) reference to Bufo coccifcr in his
discussion of the evolution of North American bufonieis
is unclear because he does not give the provenance of
the toads he analyzed. Lynch and Smith (1966) referred
specimens from the Tehuantepec region of Oaxaca, Mexico
to B. ci/clndcii; however, this designation was not followed
by nn)st subsequent authors (Porter, 1967). Toads from the
Tehuantepec region are distinct and aiiopatric, with respect
to the toads we refer to B. cyclndcii on tlu' slopes ol tiie Sierra
Madre del Sur in Guerrero and western Oaxaca.
There are no records of Bufo coccifcr from the Pacific
Coastal Plain of Chiapas, Mexico. Despite tiie lack oi
records in this region, we note that much of the region
seems to be suitable subiiumid habitat. Subhumid forest
and scrub habitats extend along the coastal plain here, but
grade into more humici forests along the base of Volcan
Tacanci (Johnson, 1989; Rzedowski, 1994). Although the
lone specimen from Tapachula appears to have been
found in a humid region, we note that the historical humid
forests (W. E. [3uellman, pers. comm.) along the coastal
margin of Volcan Tacana have been completely converted
to crop and pasture lands (pers. obs.). Further fieldwork
along the Pacific Coastal Plain of Chiapas is warranted
to \eiif\ the allopatr\' of the population of B. coccifcr in
the Islhnnis ot Ti'huanatepec, with respect to conspecifics
in Ciuatemala. Johnson (19S9) implied that B. coccifcr
occurs along the Pacific Coastal Plain of Chiapas, perhaps
based on records from Soconiisco, Chiapas, that were
Toads oi nil-; Brio C(kxii ii< C()m1'L[:.\
11
[■ij;. li- \Lip of Central AmcriLM showing the geogr.iphie distribution of Bufo coccifcr. B. ibarnii .ind B. poi Ifi i.
listed by Porter (1963); these records actually represent
misidentified specimens referable to 6. imlliccpf. The single
specimen from Tapachula, Chiapas (MVZ 177594), is a
recently metamorphosed specirnen tiiat originally was
misidentified as B. luetkeiii. This record seems to be the
source of information that resulted in B. luetkenibis'mg listed
incorrectly as part of the Mexican herpetofauna (Johnson,
1989; Flores-Villela, 1993)— B. luctkcni does not occur in
Mexico. Johnson (1990) correctly stated that records of B.
coccifcr from the Grijalva Valley of Chiapas, Mexico, were
based on misidentified specimens.
Biifo ci/clndcii Lynch and Smith
Figs. 3-5
Bufo cydaden Lynch and Smith, 1966:19. Holotype: UIMNH 57142.
Type loCiiHty; 3 mi |4.8 km] S Putio de Guerrero, Oaxaea, Mexico.
Bufo Li/cliidcii — Porter, 19h7 (decKired luvucii (luluuiii\; Frost, 1985 |in
part]; Frost, 2003 |in part].
Bufo aicn/I-i— Smith and Ta\ lor, I94S |in part]; Porter, 196,3, 1965 |in
part]; Villa et al.. 19SS |in part): Flores-Villela, 1993 |in part); Campbell,
1999 [in part].
Diagnosis. — A small species of Bufo (males to 54
mm SVL; females to 62 mm SVL), having the following
combinationofcharacters:(l)tympanumevidentexternally,
about 40-50% diameter of orbit in males, about 40-45%
in females; (2) canthal, supraorbital, supratvmpanic,
postorbital, preorbital, pretympanic, and supralabial
crests present; (3) cranial crests well developed, robust,
except parietal crest thin, low, or absent; (4) tibia short,
about 34% SVL; (5) feet short, about 34 % SVL; (6) dorsal
tubercles medium to large-sized, elevated, rounded,
densely arranged on middorsal region, becoming large
12
SciENTiiic Pai'i:rs. Naturai History Mi siim.The UNiviRsin oi Kansas
and conspicuously spinosc Litciallv; (7) ventral tubercles
reiati\el\' large, granular, with conspicuous spinose
apices; (8) lateral descending row of enlarged tubercles
absent; (9) skin texture not sexually dimorphic; (10)
vocal slit unilateral in male; (11) iii. intcrln/oidciifi poorly
differentiated from )/;. intcniiaiidibuhnis, but differentiated
posteric^rly forming a large, unilobed vocal sac with
heavy black pigmentation; (12) snout shape rounded in
lateral view, pointed in dorsal view; (13) parotoid glands
round to ovoid and large, about 1.5-2 times size of eyelid;
(14) skin between cranial crests on top of head with few
to many scattered, low, rounded tubercles; (15) ventral
cciloration whitish cream, with scattereci dark brown,
indistinct to distinct, punctuations in some individuals.
Biifo ci/tindcii may be distinguished from all other
members of the Bnfo coccifcr Group by the combination of:
its relatively small size; relatively large, scattered, rounded
tubercles on the middorsum; large, spinose tubercles on
the dorsolateral and lateral surfaces; relatively large,
spinose tubercles on the venter; large parotoid glands;
weakly developed (or absent) parietal crest; and presence
of scattered, dark brown punctations on the venter (but
this last character is variable amtmg specimens).
Biifo ci/cladcn differs from B. coccifer by being smaller
(males of B. coccifcr to 62 mm SVL, females to 82 mm),
and possessing the following characters: large, elevated,
densely arranged middorsal tubercles (middorsal tuber-
cles relatively small, scattered in B. coccifcr); large,
strongly spinose dorsolateral and lateral tubercles
(dorsolateral and lateral tubercles smaller in B. coccifcr,
variably roundeti or moderately spinose); relatively large,
strongly spinose ventral tubercles (ventral tubercles small,
rounded, or \sith tiny spinose apices in B. coccifcr); weakly
de\'eloped, or no, parietal crests (parietal crest relativeK'
well developed in most specimens of B. coccifcr); and an
adverti.sement call with a higher pulse rate (Table 1, Fig.
2). Biifo cycladcn differs from B. pisinnus by having: much
larger, distinctly spinose dt)rsolateral tubercles (smallei-,
more rounded in B. pisinnus) and relatively larger
parotoid glands (1-1.5 times si/e of eyelid in B. pisinnus).
Superficially, B. cyclndcn resembles B. signifcr, but it is
smaller (males of B. si^nifcr to 64 mm SVL, females to
77 mm) and has the following features: a whitish-cream
venter, with or vvitht)ut scattered dark brown punctations
(venter with bold brown-black marbling in /'. sis^nifcr);
and smaller and less spinose tubercles overall (but this
latter character is somewhat variable and subjective).
Bufo cyclndoi differs from B. ibarrai bv: being smaller
(males of B. ibarrai to 82 mm SVL, females to 94 mm);
having males with conspicuously spinose dorsolateral
and lateral tubercles; having relali\el\' larger, more
densely arranged middorsal tubercles (all such tubi'rcles
relatixely small, scattered, rounded in males of B. ibarrai);
and having rounded middorsal tubercles in females
(tubercles spinose in females of B. ibarrai). Bufo ci/cladoi
differs from B. portcri by: having sharply spinose lateral
tubercles in males (rounded in males of B. portcri); and
relatively thin supratympanic crests (large, bulbous in B.
portcri). Morphometric \ariation is summarized in Table
Distribution and Ecology. — Bufo cycladcn occurs in
a relati\eh' narrow ele\ational band along the Pacific
slope of the Sierra Madre del Sur in Cjuerrero and Oaxaca,
Mexico (Fig. 7). Most specimens ha\e been collected at,
or near, Agua del Obispo, Guerrero. Althc)ugh ctimmcinly
cited in the literature, Agua del Obispo does not appear
on most maps because it is a private hacienda; it is located
near the 1000-m contour (17°18' N, 99" 28' W) along the
o\d high\s'a\' between the towns of Tierra Colorada and
Chilpancingo, Guerrero. One other series (including
the holotype, UIMNH 57142) is known from Putia de
Guerrero, Oaxaca; this locality lies at an elevation similar to
that of Agua del Obispo, Guerrero, about 150 km (airline)
eastward on the same slope of the Sierra Madre del Sur.
100°
1
V
k. 20°
▲ B.cycladen ^~-~~.^_^^*-
• B. pisinnus ^"^^-—^
1 < 300 m
1 300-900 m
1 1 900-2100 m
^^° WM =• 2100 m
0 100
15°
Kilometers
100°
I ij;. 7. Map ol LiMitiMl Mexico with dot localities tor Bufo cudadcii
iTid K. pii^iiiiiu!^ (see species account lx>lou) indicating specimens
■x.iTiiined, which reprc-enls all known Imalities.
Toads oi- thk Blfo coccii i:k Complex
13
Presumnblv, the distribution of this species is continuous
along the middle-elevations of this slope, between these
two localities. Campbell and Duellman (2000:19) described
the habitat at Agua del Obispo as "...an area of scattered,
small pines and brush that appears to be ecotonal between
tropical deciduous forest, which is found slightly below,
and pine-oak forest, which is present in the mountains
above." Field work in this region by JRM and associates
in 2000 and 2002 reinforced that B. ci/chulcii seems to be
restricted to this narrow, ecotonal habitat. In this area, we
encountered: B. Jiuiriinn^ and B. iiuiniioivtis at the lowest
elevations between Acapulco and near Las Cruces; only
B. manuorcits in the vicinity of Tierra Colorada (±M00 m);
B. iimriinif and B. cycladcn near Agua del Obispo; only B.
l>crplcxii> in tine Zumpango del Rio/Chichihualco area;
and only B. occidciitalis at high elevations (over 2000 m)
near Carrizal de Bravo. Da\'is and Dixon (mtiS) reported
some additional records of Bufo from along this transect,
but we have not verified the identifications of those
specimens. The sporadic appearance of B. iiuirinii^ along
this transect is noteworthv; we also note that Davis and
Dixon (1965) did not report this species from anywhere in
the area.
Tadpole. — The tadpole of B. ci/chnicii is unknown, as
are aspects of its reproductive biology. Davis and Dixon
(1965) reported females collected on 22 June " . . .contained
numerous small eggs."
Remarks. — The type locality for this taxon is near
Putla, Oaxaca, Mexico, which is located at about 750 m
elevation on the Pacific slope of the Sierra Madre del Sur;
specimens designated as paratypes were collected at the
type locality and from around Agua del Obispt^, Guerrero,
Mexico (Lynch and Smith, 1966). However, these authors
referred all Mexican populations of B. coccifcr to this new
taxon (thereby including populations of three different
species of toads, which we have identified as B. coccifci;
B. cycladcu, and B. yifiiniiis). Furthermore, it is evident
from their map (Lyncli and Smith, 1966:fig. 2) that their
comparative samples of "B. coccifcr" from Guatemala,
Honduras, and Nicaragua may have included specimens
that we recognize as B. coccifcr, B. iharrai, and B. portcri.
Owing to this confusion, the diagnosis of B. cycladcn
presented in the original description has not been widely
accepted (Porter, 1967; McDiarmid and Foster, 1981;
Frost, 1985, 2003). We apply the taxon 6. c\/clndeii only to
those populations on the Pacific slope of the Sierra Madre
del Sur of Guerrero and Oaxaca, Mexico. It seems likely
that the confusion associated with 6. cyclnden is the result
of authors' having inadvertently compared a variety of
different species in their efforts to distinguish B. coccifcr
and B. ciicladcii. Mendelson (2001) provided additional
comments on this problem.
Bufo iharrai Stuart
' Figs. 3, 8, 9
Hufo iharrai Stuart: I S9. Holotypc: UMM7. 108000. Type kicality:
.AsiTiMdero San Lorenzo |about 12 airline km sli^hth' East of North of
LiLipa], Jalapa, GuatemaLi, 1725 ni.
Bufo coccifcr — Cope, 1887 |in part]; Meyer and Wilson, 1971 |in part];
Campbell and Vannini, 1989 |in part]; Campbell, 1999 |in part]; McCranie
and Wilson, 2002 [in part].
Hufo iharrai — Lyndn and Fugler, 1965 |in part]: Porter, 1966; I'rost,
19S5; Camplx'll, 1999; Mendelson, 2001; Frost, 200.1.
Bufo luicrotif — Sdimidt and Stuart, 1941 |in part, tor reference to
UMMZ840H3].
Diagnosis. — A large species of Bufo (males to
82.4 mm SVL; females to 94.4 mm SVL), having the
following combination of characters: (1) tympanum
e\'ident externally, about 45'7- diameter of orbit in
males, about 40' r, in females; (2) canthal, supraorbital,
supratympanic, postorbital, preorbital, pret\-mpanic.
Fig. 8. Lateral aspects ot the heads ot adult male specimens of Bufo
iharrai (upper: KU 58413) and B. pi^iiiuiis (lower: UMMZ WED 233723;
holot\ pe). Scale bar = 1 cm.
14
Scientific Papers, Natural History Museum, The University of Kansas
Fig. y. Dorsjl and vi-ntrdl dspects of representative adult specimens of Bufo ibiinni from Departamento Guatemala, Guatemala (male, left: UTA
A-25825, SVL = 69.S mm; female, right: UTA A-47572, SVL = 93.3 mm) and B. pi^imnis from Michoaain, Mexico (male, left: UMM2 115353 [WED
10971], SVL = 43.5 mm; female, right: UMMZ 121578, SVL = 60.4 mm).
parietal, and supralabial crests present; (3) cranial crests
well developed, parietal crests low, thin, sharply angled
medially; supratympanic crest large, bulbous; (4) tibia
short, about 40% SVL; (5) feet short, abi>ut 40';, SVL; (6)
middorsal tubercles sparse, roundecl, becoming spinose
lateralh' in females, all dorsal tubercles in males rounded,
usually indistinct or absent lateralh'; (7) wntral tubei-cles
areolate, non-spinose in males and finelv spinose in
females; (8) lateral descending row of enlarged tubercles
indistinct or absent; (9) skin textinv sexualK- dimoiphic;
(10) vocal slit unilateral in male; (11) ni. iiilcr!n/oulru>
poorly differentiated from ;;;. iiilciiiuiiKlibitliiri^. but
differentiated posteriorly, forming a large, unilobed vocal
sac with variable amounts of pigmentation; (12) snout
shape rounded in lateral \'ievv, weakK' pointed in dorsal
view; (13) parotoid glands large, ovoitf, length about
2 times size of eyelid; (14) skin between cranial crests
usually smooth, lacking tubercles; (15) \'entral coloration
dull cream with some diffuse gra\' mottling in some
inctividuals.
Biifo ibarrai is the largest species in the B. coccifer Croup
and differs from all species except B. portcri bv: ha\'ing
i"elati\el\' massi\e cranial crests, with the supratympanic
crest being distinctb' bulbous; and ha\ing sexually
dimorphic skin texture with distincth' roimded lateral
tubercles in males and spinose lateral tubercles in females.
Bufo ihiiiiiii closely resembles B. porlcvi but differs b\: being
larger (males to 82.4 mm SVL and females to <-)4.4 mm SVL
\s. S^.M mm in males and 76.2 mm in females); and ha\ing
the caudal muscuLiture ot the tadpole uniloini pale blown
(cream and hea\il\' punctated with brown in B. portcri).
Morphometric \ariation is summarized in Table 2.
Distribution and ecology. — Bufo ihnrrni occurs at
moderate ele\ations (1360-|9S() m) in the highlands of
Toads of the Blfo coccii kk Compi.bx
15
Fig. 10. Map of Guiiteniiila and western Honduras with dot
localities for Bufo ibnrnii and B. poilcn (see species account below)
indicating specimens examined.
central and southern duatemala (Mendelson, 2001), and
new information presented here extends that range into
contiguous regions of Honduras (Fig. 10). The Honduran
localities range up to 2020m elevation, and all represent
Premontane Moist Forest (e.g., McCranie and Wilson, 2001:
pl.2E) or Lower Montane Moist Forest (e.g., McCranie
and Wilson, 2001:pl.4E) and lie in the western ranges of
the Southern Cordillera Region (sensu McCranie and
Wilson, 2001), in the departments of Intibuca, Lempira, and
Ocotepeque (Fig. 10). These discoveries refute Mendelson's
(2001 ) premature speculation that B. ibnvrni ma\' be endemic
to Guatemala and refute McCranie and Wilson's (2001)
conclusion that B. ibanai does not occur in Honduras.
Tadpoles. — The tadpole was described bv Mendelson
(2001 ).
Remarks. — Altliough Mendelson (2001) discussed
diagnostic features that distinguish B. coccifcr and B. iharrai,
he did not present multi\ariate analyses of morphometrv.
In addition, Mendelson's (2001) research was based only
on specimens from Guatemala. During the course of this
stud\', Eric N. Smith kindly sent to us specimens of B.
iharrai he collected in Ocotepeque, Honduras. These toads
are consistent with the diagnosis presented b\ Mendelson
(200 1 ) and have a similar (0.67' <'. di\-ergent) mitochondrial-
DNA sequence to B. iharrai collected at the opposite end of
the range of the species, in El Quiche, Guatemala. We also
discovered additional specimens from western Honduras
among museum collections (Appendix II) that match tiie
diagnosis provided b\' Mendelson (2001 ); these specimens
Bufo pisinuus new species
Figs. 3, 8, 9
Bufo coccifcr Porter, 1%3 [in part]; Frost, 1985 |in part]; Villa et al.,
mSS |in part]: Flores-Villela, IW.I |in part]: Campbell, 1999 |in part];
I i-ost, 2003 |in part],
Bufo ci/clihicn Lvnch and Smith, l"-l(i(i |in part, tor reference to
specimens from Michoacan, Mexico].
Holotype.— UMMZ 233723 (WED 10973), an adult
male from 6.2 mi [10 km] E Apatzingan, 1100 feet [335 m]
elevation, obtained b\' W. E. Duellman and R. E. Etheridge
on 2 August 1936.
Paratypes. — All from Michoacan, Mexico: 6.2 mi [10
km] E Apatzingan, 1100 ft [333 m] (UMMZ 115353 [15
specimens]); 7 mi [11.2 km[ E Apatzingan, 1100 ft [335 m]
(UMMZ 112794 [6 specimens), 115335); 1 mi [1.6 km[ W
Apatzingan, 1100 ft [335 m] (UMMZ 115354); 3 mi [4.8
km] S Lombardia (UMMZ 121578).
Diagnosis. — A small species of Bufo (males to 51
mm SVL; females to 62 mm SVL), having the following
combination of characters: (1) tympanum evident ,
about 35^0 '}o diameter of orbit in males, about 40-50%
in females; (2) canthal, supraorbital, supratympanic,
postorbital, preorbital, pretympanic, supralabial crests
present, parietal crest a thin ridge, or absent; (3) cranial
crests weakly developed, thin, low; (4) tibia short, about
34';;, SVL; (5) feet short, about 36% SVL; (6) dorsal
tubercles small to medium sized, low, rounded, relatively
densely arranged middorsally, becoming smaller and
spinose laterally; (7) ventral tubercles tiny, evenly
granular, appearing smooth, especially in males, but tiny,
spinose apices apparent under microscope; (8) lateral
descending row of enlarged tubercles absent; (9) skin
texture not sexually dimorphic; (10) vocal slit unilateral
in male; (11) ;;;. iiitcrln/oideus poorly differentiated from ;;/.
iutcniiaiidihiilaris, but differentiated posteriorly forming a
large, unilobed vocal sac with heavy black pigmentation;
(12) snout shape acutely rounded in lateral view, sharply
pointed in dorsal view, snout shape in females similar but
more genth' rounded in lateral view; (13) parotoid glands
ovoid, about 1.0-1.5 times size of eyelid; (14) skin between
cranial crests on top of head usually with many scattered,
low, rounded tubercles; (15) ventral coloration usually
16
Scii-Miiic Paplrs, Nailkal History Museum, Thh Umvl;rsha oi Kansas
immaculate whitish cream, some indixiduals with tinv,
scattered black flecks.
Biifo pnsiiiiitif can be distinguished from all other
members of the B. coccifcr Group b\' its small si/e, and
possession of: relatively weakly developed cranial crests,
the parietal crest appearing only as a thin ridge among
surrounding tubercles and frequentlv is absent; smaller,
more densely arranged dorsal tubercles; and smaller,
less spinose ventral tubercles. Bitfo pisinniis differs from
B. coccifcr by having: a relatively shorter snout and an
advertisement call with a higher pulse rate (120 pulses
per sec vs. maximum of 115 pulses; Table 1, Fig. 2).
Biifo pisi)uni^ differs from B. cycladcn by having: much
smaller, and less spinose dorsolateral tubercles (large
and conspicuously spinose in B. cycladcn); and smaller
parotoid glands (about 1.5-2.0 times size of eyelid in B.
cyclndcn). Biifo p'ifiiiniis differs from B. sigiiifcr by having:
a whitish-cream venter, with or without scattered black
flecks (venter with bold brown-black marbling in B.
sigiiifcr); smaller parotoid glands (about twice size of
eyelid in B. sigiiifer); and an advertisement call with a
higher frequency and faster pulse rate (Table I, Fig. 2).
Bufo pisiimiis differs from B. ibarrai by having: smaller,
more rounded parotoid glands (glands in B. ibnrnii ovoid,
much higher, and about twice size of the eyelid); and
rounded mid-dorsal tubercles in both sexes (tubercles
in B. ibarrai rounded in males, spinose in females). Bufo
pisiunus differs from B. portcri by having: sharply spinose
lateral tubercles in males (rounded in males of B. portcri);
and a relatively thin supratympanic crest (large, bulbous
in 6. portcri).
Description of holotype. — Body robust; head wider
than long, width 3M.1';, SVL, length 35.3";- SVL; snout
sharply pointed in dorsal view, rounded in profile,
rostral keel distinct; canthal, preorbital, supraorbital,
pretympanic, supratvmpanic, and postorbital crests
present, distinct; parietal crests reduced, barely distinct;
skin on top of head co-ossified; nostril not protuberant,
directed dorsallv; can thus rostralis forming distinct, raised,
canthal crest; loival region cinna\'i-; lip distinct, rounded;
suborbital crest present, distinct, extending from angle of
the jaw anteriorlv to level of anterior margin ot orbit; notch
at symphysis of upper jaw pivsenl, distinct; e\'e-nostril
distance 5. 7'.r, diameterof orbit; t\-mpaniim distinct, nearh'
round; tympanic annulus distinct onl\' along anterior and
ventral margins, upper margin contacting supratympanic
crest, posterit)r margin obsciu-ed by overlying llesh.
Forelimb short, robust; hand broad, with short, slender
fingers; relative length ot fingers: II • I < IV ■ III, webbing
and lateral fringe on fingers absent; tips ot lingers not
enlarged, smooth dorsallv, demarcated proximalK' b\-
distinct dermal told; palmar tubercle elistincl, large,
ovoid; pollical tubercle smaller than palmar tubercle.
ovoid; subarticular tubercles distinct, elevated, triangular
in profile, single except distal tubercle on Finger 111 bilid;
supernumerary tubercles of unequal size, small, distinct,
scattered evenly over palm and ventral surfaces of
fingers; nuptial excrescences present as brown granular
patches on medial surfaces of Fingers I and II. I iind limbs
short, slender, tibia length 35.8',' SVL; foot length 38.0%
SVL; tarsal fold absent; outer metatarsal tubercle minute,
elevated, ovoid; inner metatarsal tubercle slightly larger
than outer metatarsal tubercle, distinctly elevated, ovoid;
toes long, slender, relative lengths of toes: I < 11 < V < III
< IV; lateral fringe present on Toes II, 111, and \', absent
on Toes I and IV; webbing thin, webbing formula 11 —
3II2— 3III2— 4IV4— 2V; tips of toes not enlarged, smooth
dorsally, demarcated proximally by distinct dermal fold;
subarticular tubercles distinct, elevated, triangular in
profile, bifid; supernumerar\' tubercles unequal in size,
distinct, distributed evenlv over \-entral surfaces of foot
and toes.
Skin on dorsum of bodv rugose with e\enl\'
distributed, small, rounded tubercles of relatively equal
size, becoming sharpiv pointed laterally; parotoid glands
about same size as evelids, o\oid, oriented perpendicular
to midline of bodv; lateral rov\' of enlarged tubercles
barely evident; dorsal surface of head smooth with few,
small, rounded tubercles scattered in interspaces between
cranial crests; dorsal surfaces of limbs covered with small,
weakly pointed tubercles; skin on throat and other ventral
surfaces granular, covered with tinv flattened and weakly
pointed tubercles.
Choanae small, rounded, widely spaced; teeth and
odontoids absent; tongue long, ovoid, about four times
as long as wide, free posteriorly for about one-fourth its
length; vocal slit unilateral, sinistral.
Coloration of holotype. — In preser\ati\e (ethanol),
dorsum ot bod\' and limbs mottled evenly with pale brown
and dark brown markings; irregular cream blotches present
posterior to each parotoid gland; top ot head uniform dark
brown with distinct cream interorbital bar; distinct cream
middorsal stripe extending from snout to posterior end
o\ urostvie, irregularlv widened at se\'eral points along
its length; lateral sin-faces pale brown with dark brown
flecking. Ventral surfaces nearh' immaculate cream: dark
\'ocal sac visible through gular skin.
Measurements of the holotype (in mm). — SVL 45.0,
HI. IS», IIW 17.(1, IL Ih.l. 14 17.1, orbit diameter 5.4,
tympanum diameter 23, suprahmpanic crest length 2.7,
parotoid gland length ^.4, pai'otoid gland width 4.8.
Coloration in life. — Duellman (l'-)6l:2I) described
coloration "...yellowish tan ground color with dark brow n
spots; middorsal stripe is deep ncIIow or ci-eam color. 1 he
\ enter is dust\- cream color, anil the iiis is pale gold."
Toads of the Bcro cocciier Complex
17
Variation. — Morphometric Vciriation among specimens
examined is summarized in Table 2. The parietal crest is
poorly developed in most specimens, and essentially
absent in a few individuals (e.g., UMMZ 121578). Some
individuals (e.g., UMMZ 112794) have a dark-brown
ground color on the dorsum, which effectively obscures
the dark brown dorsal blotches; most specimens have a
pale brown to grayish ground color on the dorsum, and the
overlying brcnvn blotches are distinct. The brown blotches
on the dorsum may be relativelv small, incorporating one
to three dorsal ttibercles (e.g., UMMZ 115353 [WED 10972]),
or the blotches may be larger, incorporating more than 12
dorsal tubercles (e.g., UMMZ 115353 [WED 10974]). The
narrow middorsal cream stripe invariably is present in all
individuals examined. Scattered black flecks on the ventral
surface may be absent {c.;^., UMMZ 121578), restricted to
the peripheral ventral surfaces (e.g., UMMZ 1 15353 [WED
10970]), or scattered relatively evenly across the venter (e.g.,
UMMZ 115354 [WED 10976]).
Etymology. — The name pisiiiiiiis is the Latin word
pi>iuiiii<, meaning small, in reference to the diminuti\'e
size of this species.
Distribution and ecology. — Bufo pisiiuiii^ is known
only from the Tepalcatepec Valley, which represents the
western extension of the Balsas Basin (Fig. 7). Duellman
(Field Notes, 16 July 1960) described the area arciund
Lombardia, Michoacan, as having grassy areas interrupted
by areas of mescjuite with many reddish rocks and barren
areas. Duellman (1961) suggested that this species is likely
widespread in region, but noted that it does not occur along
the coast of Michoacan. Duellman ( 1 961 ) reported breeding
choruses in muddy ditches and flooded grassy fields after
heavy rains in June and August.
Tadpoles. — The tadpole of B. pifiiniiiis is unknown.
Bufo porteri new species
Figs. 3, n, 12
Bufocoi'iifcr — MevtTiind Wilson, l'-)7I |in p>irt|; iTost, 198S [in portl;
McCranie dnd WiKon, 2002 |in part): Campbull, Iwm |in part]: Frcist, 20(1^
|in part|.
Bufo ibarnii — l.vnch iind Fiigler, I'-Ui? |in part].
Holotype. — KU 97519, and adult male from 6 mi
[9.6 km] NE Escuela Panamericana, Cerro Uvuca, 520(1 ft
[1584 m], Francisco Morazan, Honduras, obtained by K.
R. Porter on 30 June 1964.
Paratypes. — All from Franciso Morazan, Hondinas:
6 mi ]9.h km] NE Escuela Panamericana, Cerro Uvuca,
5200 ft [1584 m] (KU 97520-26); 6 mi 19.6 km] NE Escuela
Panamericana, Cerro Uvuca, 6000 ft 11828 m] (KU 97514);
W slope Cerro Uyuca, 1650 m (KU 103221); Parque
Nacional LaTigra aLxiveSan Juancito, 2100 m (KU 192294);
Cerro La Tigra, 1840-1890 m (KU 194216-19, 209249).
Fig. 11. Lateral aspects of the heads of adult male specimens of
Bufo portcn (upper; KU 97520) and B. ngnifcr (lower: AMNH 69626).
Scale bar = 1 cm.
Diagnosis. — A small to medium-sized species of
Blip (males to 59.9 mm SVL; females to 76.2 mm SVL),
having the following combination of characters: (1)
tympanum evident externallv, about 40',, diameter of
orbit in both males and females; (2) canthal, supraorbital,
suprat\'mpanic, postorbital, preorbital, pret\'mpanic,
parietal, and supralabial crests present; (3) cranial crests
well developed, parietal crests low, thin, supratympanic
crest large, bulbous; (4) tibia short, about 37' ,'. SVL; 5) feet
short, about 37"^. SVL; (6) middorsal tubercles sparsely
arranged, rounded, becoming spinose laterallv in females,
all dorsal tubercles in males rounded, usually bect)ming
indistinct or absent laterallv; (7) ventral tubercles areolate,
non-spinose in males, fineh' spinose in females; (8) lateral
descending row of enlarged tubercles indistinct or absent;
(9) skin textu re sexuall\'dimorphic;( 10 )\ocal slit unilateral
in male; (11) iii. iiitciin/ouicu> poorly differentiated from
III. iiitcniiiiiidibiilnrifi differentiated posteriorly, forming a
Scientific Paphrs, Naturai. Hisiory Museum. The University of Kansas
Fig. 12. norsjj iind ventral aspects ot representative adult specimens of Biifo portcrl from Francisco Morazan, Honduras (male, left: KL' ^7519,
SVL = 51.6 mm; female, right: KU y7.S14, SVL = 60.7 mm) and B. signifcr Panama (adult male, left: TNHC 31341, SVl, = 49.7 mm; subadult female,
right: KU 1 1 3359, SVI . = 54. 1 mm ).
Itirgc, unilobed vocal sac hcavih' pii^LMiiented in black;
(12) snout shape roiindccl in lateral \ic\v, wcakh' poifited
in dorsal xiew; (13) pafotoid glatids moderate, rounci to
ovoid, length about 1.00-1.23 titnes size of eyelid; (14)skifi
between cranial crests usually smooth, lacking tubercles;
(15) ventral coloration is dull cream with some diffuse
gray mottling in some individuals.
Biifo poiicii ma\' be distingiushed trotn all other
tnember of file B. coaifcr Group, except B. ihninu, the
sexual dimorphism in the texture of the dorsal skin
and the relatively robust cratiial crests, especialh' the
supratympanic crest. Biifo poiicri closely resetnbles />'.
ibanni but differs by being smaller (tnales to 54.9 mm
SVL and females to 76.2 mm SVL \'s. males to >S2.4 mm
SVL and females to 94.4 mtii SVL); adult male B. ]wrtcn as
small as 43.9 mm SVL have been observed (e.g., LSUMZ
46431 ). The caudal musculature of the tadpole of B. poiicri
is creafTi w ith hea\ \ brown punctations (See description
b\' McCranie and Wilson, 2002:173; caudal musculature
if! B. ibarrai is unifortii pale browti.)
Description o( holotype. — Bod\ robust; head \\ ider
than long, width 40.0% SVL, length 36.4% SVL; snout
sharply pointed in dorsal view, pointed in profile, rostral
keel distinct; canthal, preorbital, supraorbital, pretytnpanic,
supratympatiic, and postorbital crests present, distinct;
supraorbital atid suprat\'mpanic crests distintly thickened;
parietal crests present, not reduced; skin on top of head
co-ossified; nostril protuberant, directed dorsally; canthus
rostralis forming distinct, raised, canthal crest; loreal region
concaM'; lip distinct, rounded; suborbital crest present,
distinct, extending from angle of the jaw anteriorly to le\'el
of afiterior margin of orbit; notch at s\'mphvsis of upper
Toads of ihh Buio coven ik Complex
19
jaw present, distinct; e\'e-n(istril distance 37.9",- diameter of
orbit; tympanum distinct, nearly round; tympanic annulus
distinct only along anterior, xentral, and posteroventral
margins; upper margin of tympanic annulus contacting
and obscured by supratvmpanic crest. Forelimb short,
robust; hand broad with short, slender fingers; relative
length of fingers: II < I < IV < III, webbing and lateral
fringe on fingers absent; tips of fingers not enlarged,
smooth dorsallv, demarcated proximally by distinct
dermal fold; palmar tubercle distinct, large, rounded;
pollical tubercle smaller than palmar tubercle, rounded;
subarticular tubercles distinct, ele\ated, triangular in
profile, all single except distal tubercle on Finger 111 bifid;
supernumerary tubercles of unequal size, small, distinct,
scattered evenly over palm and ventral surfaces of fingers;
nuptial excrescences present as brown granular patches
on medial surfaces of Fingers l-IIl. Hind limbs relatively
long, slender, tibia length 41.7",. SVL; foot length 40.5",'.
SVL; tarsal fold absent; outer metatarsal tubercle very
small, rouncied, indistinct; inner metatarsal tubercle much
larger than outer metatarsal tubercle, distinctly elevated,
ovoid; toes long, slender, relative lengths of toes: I < II <
V < III < IV; lateral fringe barely evident on Toes III and
IV, absent on other toes; webbing thin, webbing formula
II — 3II2 — 3III2 — 4IV4 — 2V;'tips of toes not'enlarged,
smooth dorsally, demarcated proximally bv distinct dermal
fold; subarticular tubercles distinct, elevated, triangular in
profile, bifid on Toes III and IV; supernumerary tubercles
unec^ual in size, distinct, distributed evenly over ventral
surfaces of foot and toes.
Skin on dorsum of body relatively smooth with
scattered, small, rounded tubercles of unequal size, many
bearing tiny, indistinct single keratinized apices; tubercles
on lateral surfaces indistinct, roimded or ovoid, ncit
pciinted; parotoid glancis about same size as eyelids, ovoid,
oriented slightly divergent tci midline of body; distinct
lateral row of enlarged tubercles absent; dorsal surface of
head smooth \yitlT few, small, rounded tubercles scattered
in interspaces between cranial crests; dorsal surfaces of
limbs covered with small weakly pointed tubercles; skin
on throat and other ventral surfaces smooth!)' granular.
Choanae moderate in size, rounded, widely spaced;
teeth and odontoids absent; tongue long, ovoid, about four
times as long as wide, free posteriorly for about one-foiuih
its length; vocal slit unilateral, dextral.
Coloration of holotype. — In preservative (ethanol),
bod\' pale brown with indistinct, medium-brown marbled
markings diffused across middorsal and lateral areas;
distinct, thin cream middorsal stripe present; two oblong
dull cream patches on dorsolateral areas; distinct dark
brown bar extending across area between parietal crests
and another covering area between canthal crests; all
limbs with indistinct medium brown crossbars; venter
immaculate dull cream; dark vocal sac clearly \'isible
through gular skin.
Measurements of the holotype (in mm). — SVL 51.3,
HL 18.7, HW 20.5, TL 21.4, FL 20.8, orbit diameter 7.6,
tympanum diameter 3.0, supratympanic crest length 3.1,
parotoid gland length 6.9, parotoid gland width 4.8.
Coloration in life. — McCranie and Wilson (2002:pl
'■'D) provided a color photograph of this speceis; see also
Figure 3.
Variation. — Morphometric variation among specimens
examined is summarized in Table 2. The parietal crests
in females generally are well developed and distinct;
however, the condition of the crest varies among males
and may be relatively robust (e.g., KU 209249) or reduced
to an indistinct, thin sliver of raised bone (e.g., KU 97523).
The brown dorsal blotches may be very dark brown, and
therefore quite distinct (e.g., LSUMZ 46398), or they may
be only slightly darker than the brown ground cc^lor of
the dorsum (e.g., LSUMZ 46445). The ventral coloration
is either uniform dull cream (e.g., KU 103221) or bearing
diffuse, grayish-brown marbling that ranges from
moderate (e.g., KU 97522) to extensive (e.g., KU 194216).
The middorsal cream stripe invariably is present on all
individuals examined; this stripe is quite indistinct in a
small nimiber of individuals (e.g., KU 97520) and may
appear irregular (i.e., not ft)rming a straight line; e.g.,
LSUMZ 45441).
Etymology. — The specific epithet is a patronym that
honors Kenneth R. Porter and his series of papers (e.g..
Porter, 1963, 1964, 1965)onthesystematicsof Mesoamerican
Biifo, and also recognizes his numerous field efforts that
resulted in many of specimens referred to herein.
Distribution and ecology. — Bitfo povtcvi is known
from the Hondiu-an departments of Comavagua, Francisco
Morazan, and La Paz (Fig. 10). The known localities for this
species generally represent Lower Montane Moist Forest
habitats (e.g., McCranie and Wilson, 2001 :pl. 4C) in the
Montanas de Comayagua region. These localities differ
markedly from the lower elevation, Dr\' and Arid Forest
habitats (e.g., McCranie and Wilson, 2001 :pl.lC) occupied
b\' B. coccifer.
Tadpoles. — Tadpoles leferrable to B. yoiicri (based
on geography) were described bv McCranie and Wilson
(2002:173). The tadpole of this species resembles that of B.
coccifer (McDiarmid and Foster, 1981; Sa\age, 2002), from
which it is distinguished by having submarginal papillae
on the oral disc (absent in 6. coccifer) and cream-colored
caudal musculature with brown punctations (boldly
marked with brown saddles in B. coccifer).
Remarks. — We have allocated many Honduran
specimens that previously were referred to B. coccifer
20
Scientific Paphrs, NATiiRAi. Hisiok^ Mu.shum,Thh UNivi-.Rsrn oi Kansas
(e.g., McCranie and Wilson, 2001) to the new taxon B.
porteri, or to B. ibarrai (discussed above, and Appendix II).
The difficulty of identifying specimens from this country
is exacerbated bv the resemblance of B. porteri and B.
ibarrai. The wide range of variation among specimens of
"B. coLxifcr" from Honduras described by McCranie and
Wilson (2001 ) seems to be attributable to the fact that three
relative!)' similar species occur in close proximit)' in that
country.
At a general le\el, this species appears to be
parapatric with respect to the distribution of Bnfo coccifcr.
Thus, the distribution and habitat associations of this
species, with respect to those of B. coccifer, resemble the
relationship between B. coccifcr and B. ibarrai in Guatemala
as described by Mendelson (2001 ). Inasmuch as species
of Bnfo frec]uentl\- are interfertile (Blair, 1963; Masta et al.,
2002), it is possible that B. porteri may hybridize with B.
coccifer if the species co-occur on the lower slopes of the
Pacific Versant of Honduras. Similarly, hvbrids between B.
porteri and 6. ibarrai eventually may be found.
Bnfo sigtiifer new species
Figs. 3, 11, 12
Bufo coccifer Dunn, 1933; Zweifel, 1965; Frost, 1985 [in part, tor
reference to specimens from Panama]; Villa et al., 1988 |in part, for
reference to records from Panama]; Campbell, 1999 |in part].
Holotype.— AMNH 69626, an adult male from 7 mi N
[11.2 km] and 2 mi [3.2 km] W of David, Chiriqui', Panama,
obtained bv R. G. Zweifel on 25 June 1962.
Paratypes. — All from Panama. Chiriquf: 7 mi [ 1 1 .2 km[
E ConcepcicSn (AMNH 69627); 2.5 mi [4.0 km| NE David
(TNHC 3134-43); 23 km NNE San Felix, 900 m (USNM
297511-21). Code: El Valle de Anton (AMNH 59634, 59637);
16 km S, 9 km W Penonome, 30 m (KU 11 5359-6 1 ); 3.2 km
W Agua Dulce, 15 m (KU 115362). Herrera: Jacinto, 2250 ft
[686 m] (ANSP 22341-44); 3 mi [4.8 km] SW Pan American
Hwy, on road past Potuga (UMMZ 167373). Veraguas: 14
km NE Sona, 75 m (KU 95432).
Diagnosis. — A medium-sized species of Bufo
(males to 64 mm SVL; females to 77 mm SVL), ha\ ing
the following combination of characters: (1) tympanum
evident externally, about 40-45'('. diameter of orbit in
males, about 40-50% in females; (2) canthal, supraorbital,
supratympanic, postorbital, preorbital, pret\nipanic,
parietal and supralabial crests present; (3) most cranial
crests distinct and thick, except parietal crests low, thin,
sometimes intermittent; (4) tibia short, about 35'f' SVL; (5)
feet short, about 36' V, SVF; (6) dorsal tubercles medium-
sized, prominent, roimded, rel,ili\el\' denseh' arranged
middorsally, becoming smaller, more concentrated, antl
spinose laterally; (7) ventral tubercles granular, willi
small, distinct spinose apices; (8) lateral desci'nding row
of enlargeti tubercles absent; (9) skin texture not sexualh'
dimorphic; (10) vocal slit unilateral in male; (11) iii.
iiiterh\/oiiieiis poorly differentiated from iii. iiiteriiniiulibulari^,
but differentiated posteriori)' forming a large, unilobed
vocal sac with heavy black pigmentation; (12) snout
shape rounded in lateral view, pointecf in dorsal \'iew;
(13) parotoid glands round to subovoid, about twice size
of e\'elid; (14) skin between cranial crests on top of head
usually with few, scattered, low, rouncfed tubercles; (15)
ventral coloration cream with distinct, bold, brown-black
marbled pattern, becoming indistinct o\er pehic patch.
Bnfo ^i^'^iiifer may be distinguished from all other
members of the Bufo coccifer Group bv ha\'ing a cream
venter o\'erlain with a distinct, marbled pattern of brown-
black markings. All t)ther species in the group have
immaculate, ornearh' immaculate, cream ventral surfaces
with the exception that some indi\iduals of B. cucladeu may
ha\'e some dark brown mottling, and some indix'iduals of
B. pisinniis mav ha\'e some tiny, black flecks. Bufo si^i^uifer
differs from B. coccifer by having: relatixel)' thinner and
lower parietal crests (typically higher and thicker in B.
coccifer); relatively smaller tympana; and an advertisement
call with a lower frequencv and pulse rate (Table 1, Fig. 2).
Bufo :^igiiifer is superficially quite similar to B. ci/cladeu but
differs by: being larger (males of B. cycladen to 54 mm SVL,
females to 62 mm); having a distinctly marbleci \'entral
pattern (variably present, but always less developed in
B. cycladen); and having generallv smaller, less spinose
tubercles overall (but this latter character is someu'hat
\'ariable and subjective). Bnfo signifer differs from B.
pisinnub bybeing larger (males of B. pi^inuns to 51 mm
SVL, females to 62 mm) and by having: larger middorsal
tubercles (small in B. pisinnns); larger parotoid glands
(glands about 1.0-1.5 times size of eyelid in B. pisiniius);
spinose tubercles on the venter (granular, non-spinose in
B. pisi)i}uis); an advertisement call with a lower frequency
and pulse rate (Tablel, Fig. 2); and better-developed cranial
crests, especially with respect to the parietal crest (crests
weaklv developed, and parietal crest very reduced or
absent in B. pissiiin^). Bnfo signifer differs from B. ibarrai by
being smaller (males of B. ibarrai to 82 mm SVL, females
to '^'4 mm) and b\' having: smaller, more rounded parotoid
glands (glands in B. ibarrai ovoid, about twice the size of
the e\'elid); and rounded mid-dorsal tubercles in both
sexes (tubercles in B. ibarrai roimded in males, spinose
in females). Bnfo fiignifer differs trom B. porteri by having:
sharplv spinose lateral tubercles in males (roimded in
males of B. porteri); and a relatively thin supratympanic
crest (large, bulbous in B. porteri).
Description of holotype. — Body robust; head wider
than long, width 39.9', SVL, length 34.7';;, SVL; snout
sliarph' pointed in dorsal \'iew, rounded in profile, rostral
keel distinct; canthal, pivorbital, supraorbital, pretympanic,
siiprat\mpanic, and postorbital crests present, distinct;
Toads of the Bufo coccih.r Complex
21
parietal crests present, relati\el\' indistinct; skin on top
of head coossified; nostril protuberant, directed dorsally;
canthus rostralis forming distinct, raised, canthal crest;
loreal region concave; lip distinct, rounded; suborbital
crest present, distinct, extending from angle of the jaw
anteriorly to level midway between orbit and nostril; notch
at symphysis of upper jaw present, shallow, indistinct;
eye-nostril ciistance 47.6' r. diameter of orbit; t\mpanum
distinct, nearlv round; t\mpanic annulus distinct only
along anteroventral margin, upper margin contacting
supratympanic crest, posterior margin obscureci by
overlying flesh. Forelimb short, robust; hand broad, with
short, slender fingers; relati\'e length of fingers: II < I <
IV < III, webbing and lateral fringe on fingers absent;
tips oi fingers not enlarged, smooth dorsallv, demarcated
proximalh' by distinct dermal fold; palmar tubercle
distinct, large, round; pollical tubercle smaller than palmar
tubercle, ovoid; subarticular tubercles distinct, elevated,
triangular in profile, single except ciistal tubercle on Finger
III bifid; supernumerary tubercles of unequal size, large,
distinct, scattered evenly over palm and \entral surfaces
of fingers; nuptial excrescences present as bro\N'n granular
areas on entire dcirsal surfaces of Fingers I and II, including
lateral surfaces of tips of fingers, lateral surfaces of distal
phalange of Finger III, and on medial surface of pollical
tubercle. Hind limbs short, slender, tibia length 34.5' «'. SVL;
foot length 35.1"o SVL; tarsal fold absent; outer metatarsal
tubercle small, elevated, ovoid; inner metatarsal tubercle
slightly larger than outer metatarsal tubercle, distincth
elevated, ovoid; toes long, slender, relative lengths of toes:
I < II < V < III < IV; lateral fringe on toes barely evident
on Toes II and III, absent on other toes; webbing thin,
webbing formula II — 27,111'/, — 3III2 — 4IV4 — 2V;
tips of toes not enlarged, smooth dorsally, demarcated
proximally by distinct dermal fold; subarticular tubercles
distinct, elevated, triangular in profile, single except two
most distaltubercles on Finger IV and most distal tubercle
on Finger V bifid; supernumerary tubercles unequal in
size, distinct, distributed evenly over ventral surfaces of
foot and toes.
Skin on dorsum of body rugose with scattered,
rounded tubercles of unequal size, becoming sharply
pointed laterally; parotoid glands about same size as
eyelids, slightly ovoid, oriented parallel to midline of
body; lateral row of enlarged tubercles barely evident;
dorsal surface of head smooth with few, small, rounded
tubercles scattereti in interspaces between cranial crests;
dorsal surfaces of limbs covered with small to large, mostly
pointed tubercles; all ventral surfaces rough, co\'ered with
small, conical tubercles.
Choanae small, ovoici, widely spaced; teeth and
odontoids absent, but ventral surface of neopalatine
appears serrate; tongue long, ovoid, about four times
as long as wide, free posteriorly for about one-fourth its
length; \'ocal slit unilateral, dextral.
Coloration of holotype. — In preservative (ethanol),
dorsal areas of body and limbs mottled evenly with pale
brown and dark brown markings, with a bilateral series of
small, oblong cream markings dorsolaterally; top of head
uniform dark brown with distinct cream interorbital bar;
distinct cream middorsal stripe extending from snout to
posterior end of urostyle; lateral surfaces dull brown with
some dull cream supralabial spots. Ventral surfaces dull
cream with indistinct pale gray markings, forming loosely
reticulate pattern; dark vocal sac barely visible through
gular skin.
Measurements of the holotype (in mm). — SVL 57.1,
HL 20.5, HVV 23.0, TL l'-).9, FL 20.0, orbit diameter 8.2,
tympanum diameter 4.4, supratympanic crest length 3.4,
parotoid gland length 6.8, parotoid gland width 5.9.
Coloration in life. — Original field notes by R. G.
Zweifel { 1 2 June 1 962) describe AMNH 69625 as " . . .a rather
black and white looking toad, the whiteness coming from
80°
10°
B. agnifer
< 300 m
300-900 m
900-2100 m
I ""H > 2100 m
0 100
Kilometers
80=
Fig. 13. Map of western Panama with dot localities for Biifo >iguiffr
indicating specimens examined, which represent all known localities.
97
SciHNTiHC Paphrs. Natural History Museum, The University of Kansas
the lateral stripe which begins at tlie parotid and runs back
to the groin and by the central vertebral line. The blackness
is supplied by the dorsal blotches which are almost jet-
black. The ground color between the blotches is dark gray.
The side below the lateral line is a mixtiuv ot black and
gray. The eyelids are a stripe with grayish white and black.
The interorbital area is black with a grayish white band.
The \'entral surfaces are mottleci gray and white; there is
a dark gray spot in the middle posterior gular region. The
hind legs are barred very with dark gray and grayish white
as are the limbs right out to the tips of the toes."
Variation. — Morphometric variation among specimens
examined is summarized in Table 2. The parietal crests in
males may be relatively distinct and well formed (e.g.,
TNHC 31343, or they may be essentially absent (e.g., TNHC
31342); this crest is always relatively distinct in females.
The dorsal pattern usually consists of a contrasting array
of dark brown or black blotches over a pale brown ground
color, but some specimens (e.g., TNHC 46261 ) have a nearly
uniform, dull brown dorsimi. Nearly all specimens of
this species bear a distinctive, highly contrasting marbled
pattern on the venter; this pattern is present, but less
distinct in a few specimens (e.g., TNHC 46261, USNM
297515, AMNH 69626). The middorsal cream stripe is
present in all individuals, but may be incomplete (e.g.,
AMNH 69627).
Etymology. — The name sij^iiifcr Latin, meaning bearing
marks, refers to the ventral coloration of this species.
Reproductive biology. — The tadpole of B. signifcr is
unknown. Zweifel (personal communication) found calling
males in a "weedy, shallow, muddy roadside pool" cin 25
June 1962; also found in this pool were ElnchiitocU'is sp.
and Lcptodncti/liib in^ulnniiii.
Distribution and ecology. — This species is known
from each of the disjunct areas of Tropical Dry Forest
(Campbell and Lamar, 2004:47, color map 6) along the
Pacific Coast of Panama (Fig. 13). These regions are located
in the vicinity of city of David, and the Province of Veraguas
eastward toward the Canal Zone.
MORPHOMETRIC ANALYSES
Overall morphometric variation fc)r all species is
summarized in Table 2. Two Principal Components
Analyses were conducted: one with log-transformed data,
and one using residuals of each variable regressed on SVL.
These analyses produced similar results (not presented
here) wherein the first PC had the greatest eigenvalue
(and accounted for the majority of the variation explained )
and represented an overall size axis. Despite reasonably
high loadings for variables such as TYMP and PARW on
the second PC, plots individual scores for each specimen
did not clearly distinguish among the species. As could be
expected, Biifo coccifer showed the most overall variation,
while the smallest species (6. pisiiniiifi) was somewhat
distinct from the largest species (B. ihnrrai) along the
size axis (PCI). Considered together, the results of these
analyses indicate that, with the exception of overall size,
there has been relatively little morphometric differentiation
among the species in this group.
Variable selection procedures (forward, backward, and
stepwise) for LDA suggested retention of all nine variables.
i.e., residuals of eight variables regressed on SVL and
log-transformed SVL. The cross-validated classification-
matrix from the LDA is shown in Table 3. The majority
of individuals of all species were correctly classified.
Individuals of Biifo sigiiifer and B. porteri were misclassified
inconsistenth' among all other species. Biifo pisimiiis and
B. cycladcn had the greatest percentage (86%, 80'v,) of
correctly classified individuals, respectively. The small
size of 6. pisiiuuifi likely is responsible for this high degree
of correct classification, but we note the 11 individuals of
the quite variable species B. coccifer were misclassified as
B. pifiinius; this result suggests that small B. coccifer are
morphometrically similar tti B. pisiiniiis. Similarly, the
relatively large size of B. ibarrai likely is responsible for
the relatively high percentage (79' ,',) of correctly classified
individuals of that species (and the very few individuals of
other species that were identified as B. ibarrai). In general,
these results are consistent with the results of the PCA.
Despite its wide range of overall variation (Table 2), 77%
of indi\iduals of B. coccifer were classified correctly.
Table 3. Cross-validated classification-matrix and overall correct classification percentages (rounded to nearest integer) from Linear
Di.scriminant Analysis based on residuals of eight morphometric variables regressed on SVL and log transformed SVL from all species in the Bufo
coccifer Group; analysis includes only adult males. Values in boldface indicate number of individuals correctly classified.
Per
cent correct
N
B. coccifer
B. cycladcii
B. ibarrai
B. pi:
isiiuif
B.
sif;nifcr
B. porteri
cla
ssification
B. coccifer
106
82
4
2
11
4
3
77
B. cyctaden
30
1
24
0
2
3
0
80
B. ibarrai
79
0
2
62
0
2
3
79
B. pi'^aiiiu^
21
1
1
0
18
1
0
86
B. aignifcr
21
0
2
1
1
16
1
76
B. porleri
43
3
1
1
5
1
34
79
Toads of the Bufo coccifer Complex
23
Table 4. Voucher numbers and locality int'omiation for specimens o( Bii/o used in the molecular
analyses. GenBank accession numbers are given for I6S and cyt-b. respectively.
Taxon
Voucher No
Local itv
GenBank No.
B. ihcinai I
B. iharrai 2
B. coccifer HS
B. coccifer Nl
B. coccifer HO
B. coccifer CR
B. cycliulen
B. conifenis
B. valHceps
B. mcirimis
UT.^-ENS 10270
UTA-.IAC 19612
KU 290030
SDSNH-ARH0I3
USNM 547980
TC'WC 8399S
UTA-JRM 4607
MVZ 203775
MZFC-JRM 3868
UTA A-50864
Honduras: Ocotepeque
(iuatemala: Quiche
HI Salvador: Morazan
Nicaragua: Ometepe Is.
Honduras: Gracias A Dios
Costa Rica: San Jose
Mexico: Guerrero
Costa Rica: Cartago
Mexico; Veracruz
Guatemala: Zacapa
.A'i'927854,.'\Y927S6l
.'\Y927S55, AY927862
.•\Y927S56, AY927S63
.•\Y927857. A>'927864
.•\'i'92930I.AY929303
.^929302. AY929304
AY927858. AY927865
U52800. AY008255
AYO0821I.AYOO82I2
AY927860. AY927867
MOLECULAR ANALYSES
Sequences of 547 and 410 base pairs were obtained
for 16S and cyt-b mtDNA genes, respectively; specimen
information and GenBank Accession numbers are listed
in Table 4. These sections correspond to the 4004—1556 and
16422-16818 positions for 16S and cyt b, respectively, on the
mtDNA genome of Xeuopus [Pipidae] (Roe et al., 1985). Of
these 956 base pairs, 765 were constant, 191 were variable,
and 85 were considered parsimony-informative characters.
The partition-homogeneity test results indicated that trees
from the separate genes were not significantly different
from one another (P = 1.0). Parsimony analysis of the
16S region produced four trees. Parsimony analysis of
the cyt-b region produced one tree that was identical the
combined analysis shown in Figure 14. The differences
among the 16S trees were the monophyly/paraphyly of
Bufo iharrai, and the relationships among B. coccifer (sensu
stricto) samples. Parsimony analysis of all nucleotide data
combined evaluated a total of 2,027,025 trees, with the best
tree score of 278 steps (Fig. 1 5) and the worst tree score of
373 steps. The frequency distribution of trees scores had
a mean of 351.60 steps (sd = 14.8; g, = 1.20; g, = 0.94). The
hLRTs selected the TrN + I + G model (Tamura and Nei,
1993) as the most significant (p < 0.01; -InL = - 2612.9934)
with base frequencies of A = 0.3071, C = 0.2448, G = 0.1620,
T = 0.2860. The substitution rate matrix was: A-G = 6.4270,
C-T = 12.1788, all others were equal to 1; the proportion
6. valliceps
B. marinus
B. coniferui
B. cycladen
B. ibarrai
Guatemala
5. ibarrai
Honduras
B. coccifer
Nicaragua
6. coccifer
Honduras
S. coccifer
El Salvador
B. coccifer
Costa Rica
Fig. 14. Phylogenetic relationships among samples used in
molecular analyses; see Table 4 for specimen information. Topology
is shown from the maximum-parsimony exhaustive search. Bootstrap
values greater than 50' ». (MP /ML) are shown for each analysis above
branches and decav indices are shown below.
98/99
100/82
91
89
10
11
2
97/74
2
61
7
1
51
1
Table 5. Pair-vvisc sequence divergence of 1 6S and cyt b combmed for specimens ofBii/o used in the molecular analyses. Absolule distances are shown above
the diagonal, and Tamura-Nei corrected distances are below the diagonal.
Taxon
1
">
3
4
5
6
7
8
9
10
1 . B- il'<urrLtt 1
6
19
17
18
19
35
70
107
113
2. B. iharrai 2
0.0064
—
17
15
16
19
34
74
105
113
3. B. coccifer ES
0.0204
0.0182
—
4
3
8
40
72
106
115
4.B. coccifer m
0.0184
0.0162
0.0043
—
3
10
40
72
101
112
5. B. coccifer HO
0.0195
0.0173
0,0032
0.0032
—
9
40
74
107
116
6. S coccifer CK
0.0205
0.0205
00085
0011)7
0.0097
44
75
108
117
7. B cycladen
0.0832
0.(070
0.0440
0,0445
0,0444
0,0486
83
108
114
8. B. conifenis
0.0791
0.0841
0.0816
0,0824
0,0848
0,0855
0O950
—
106
124
9. B. valliceps
0.1261
0.1232
0.1253
0.1196
0.1277
0,1276
0.1269
0.1
235
—
109
10. B. marinus
0.1325
0.1325
0.1358
0.1332
0.1383
0. 1 38 1
0. 1 343
0.1483
0.1292
—
24
SciENTii K" Pai'i:ks, Natural Hisioky Mushum,Thi-: University oi- Kansas
of invariable sites was I = 0.5899 and the gamma shape
parameter was G = 0.4906. A ML heuristic search produced
a tree with a score of In L = -2609.8015 that was slightly
different from the MP tree. In the ML tree (not shown), the
B. ibarrai samples were basal to the remaining samples of B.
coccifer (sensu lato); however the nodes in the ML analysis
differing from the MP analysis were not supported in
the ML bootstrap analysis. The ML bootstrap values are
shown for nodes supported by more than 50% in Figure
14. Sequence divergences for the combined 16S and cyt b
sequences are shown for each sample in Table 5.
The phylogenetic hypothesis generated by our
data (Fig. 14) supports recognition of a monophyletic B.
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suggest that B. coccifer (sensu lato) represents more than one
species. Our proposed taxonomy recognizes six species in
this group, one of which (B. C}/cladc}i) was here recovered
as the sister to (B. coccifer + B. ibarrai). Knowledge of the
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26
SciENTiiic Papkrs. Natlral Histor'> ML'siaM.Tiiii Universh^ oi Kansas
APPENDIX
Specimens examined
Note that data presented here has been taken verbatim fnmi
museum catalogs and tags. We have not attempted to correct various
spellings, nor to alter original estimates of distance and elevation. We
do not consider it reasonable to re-estimate primarv locality data.
Biifo coccifer: Costa Rica: Ai .muei a: Rio CIrande (LACM 594.'$2).
PuNTARtNAS: 1.5 km W Barranca, 20 m (KU fi.S.'iSl); 4 km WNW Esparta,
45 m (KU 65397-90, 65393, 65397-98, 65400, 6.5404-05, 65414). San
Jost: Guadalupe, 1190 m (KU 65379); Esca/ii, 1100 m (KU 65380).
Guatemala: Cimqiumii a: above headijuarters of Finca San Jose, ca. 6.0
km SE Concepcion l.as Minas (UTA A-38119); Finca San Jose, ca. 6.0
km SE Concepcion Las Minas on rd to Las Presas (UTA A-381 23-26);
Concepcion Las Minas, Valle Arriba (UTA A-47566); 2.6 mi N intersection
of CA-12 and CA-10 (UTA A-38120-21); Jalaca: Volcan Jumay (UTA
A-47591). RtTALHULEU: 3.2 km N Champerico (UTA A-25814-'23); 3.7
km N Champerico (UTA A-29021-25). Santa Rosa: El Oratorio, 3.0
km E CA-8 (UTA A-38143). Hi Salvador: Chalatengo: 16.5 km WNW
Chalatengo (KU 184659-62); 10 km NE La Palma, Canton Las Pilas
(KU 184663-69), Cuscatlan: Tcnancingo, Rio Quezalapa (KU 184572
i73); 7 km W Cojutepeque, 2900 ft (KU 97495); 8 km W Cojutepeque,
2850 ft (KU 97499-504); Cojutepeque, 3220 ft (KU 97505-09); 0.5 km SE
Cojutepeque, 2520 ft (KU 97510-11); 2 km E Cojutepeque, 3250 ft (KU
97496). l.A Libtrtad: 10 mi NW Santa Tecla (KU 41411-29); 5 mi W Colon
(KU 97473-86); 5.4 mi W La Libertad, 680 ft (KU 97487-92); San Bartolo,
11 km E San Salvador, 2595 ft (KU 97493). La Union: Laguna Olomega
(KU 184574). M(1ra/an: 12 km NE Perquin, Canton, El Zancudo (KU
184578-609); grounds of Hotel Perquin Lenca, Perquin, 1150 m (KU
290030). San Salvador: 1 mi NW San Salvador (KU 42613-24); Instituto
Tropical (KU 61871-74); 16 km E San Salvador, 2880 ft (KU 97494); 3 km SE
llopango. Canton A.sino (KU 184575-77, 184670-74); San Salvador, Cuidad
Universitaria (KU 184612-58, 184715-16). Santa Ana: Rancho San Jose,
800 m (KU 65372). San Vicenit;: 1 km E El Carmen (12 km E Cojutepeque),
2620 ft (KU 97497); 6.5 km E El Carmen (17.5 km E Cojutepeque), 2700
ft (KU 97498). Honduras: Ciiolltra: 6.8 mi S Prespire, 380 ft (TNHC
31461); Choluteca (USNM 167195); 1.5 km NW Choluteca, 170 m (KU
65375-76); San Francisco de Colon, 1 130 m (KU 65374). El Paraiso: 4.4
mi SW Santa Maria, 1960 ft (KU 97512-13). Francisco Mor,-\/an: 10.3 mi
S La Vente, 530 ft (TNHC 31454); vie. of Tegucigalpa (LSUMZ 24133-34).
Ckacias A Dios: Rus Rus, 60 m (USNM 547977, 547980); Mocoron (UTA-CJF
1883-84). Olancho: 1 km NW Catacamas (LACM 47973-74, 21584-89);
Catacamas, 460 m (KU 194214); ca. Dulce Nombre de Culmi, 600 m (KU
194213). Valle: 1 mi E Goascoran (L.A.CM 47975-77); 3 mi E Goascoran
(LACM 47978-82); 5.2 mi SE Jicaro Galan, 250 ft (TNHC 31456, 31459).
Mtxico: Chiapas: road from Tapachula to Puerto Madero. Oaxaca: O.I mi
S jet 185 & 190, on 185 (LACM 38160-61 ); Juchitan (USNM 51 175); 4 mi S
Juchitcin, 120 ft (KU 974.34-42,); 1.5 mi N Juchitan, 120 ft (KU 97443); 3 mi
N Juchitan, 120 ft (KU 97444-17); 0.5 mi S Juchitan, 140 tt (KU 97448-50); 5
mi N Juchitan, 110 ft (KU 97451-62); Juchitan, 9 mi 1-; jet hvvys 190 and 1S5
(TNHC 30968); Hwy 18,5, 14.2 mi N jet Hvvy 190 ( INHC 5,3682, 53684);) mi
NW Zanatepec (TNI IC 31 .338); 20 mi W Zanatepec (TNI IC 27161 ); 3 mi W
Zanatepec (TNHC 271 65); 10 mi W Zanatepec (TNI IC 27292); Zanatepec
(TNHC 27300); 17 mi E Tehuantepec (TNHC 27282, 27284, 27286, 27288,
27290, 27305); 40 mi E Juchitan (TNHC 27291 ); edge of Tepanfepec (TNHC
2729,3,27295,29297), Nicarac;ua: Granada: shoreofl.agode Nicaragua,
ca 2 mi from Granada (LACM 67585). Man.-u.la: Managua, S shores
of Lake Managua (LACM 28165-69); Los Kobles (LACM 37957-,5S, KU
17.3951-53, 173955-.56); 2-5 km STipitapa(KU 173958), Rivas: Nandaime,
400 ft (KU 97547). Zi i aya Noktl: Leicus Creek at La Tronquera (NIPCX)
Lumber Plant), .56 mi NW Puerto Cabeza (l,At:M 14580S); Slilma Sia, 16
km SE Waspam, Comarca de El Cabo (LACM 145813).
Bufo cyclailen: Mfxico: Ck trrtro: near Palo Blanco (FMNH 99682,
99686, UIMNT I 248,34-,38); Xaltianguis (UIMNH 24833); Agua del Obispo,
KM 350-351 (UlMNl I 248,39); Agua del Obispo, KM 357 (UIMNH 24840);
4 km bevond Agua del Obispo (UIMNH 24841^5); near Agua del Obispo
(FMNH 99684-85, 99687-90, 105394, 107984-91, USNM 11548283);
Agua del Obispo (UIMNH 24846, 24848-50, 24875-76, 57143); Agua del
Obispo, 980 m (KU 86672-73); Agua del Obispo, 2900 ft (UMMZ 115357
1 6 specimens]), Oax.aca: 3 mi S Putla (UIMNH 57144-51),
Bufo ibnrrni: Guatemala: Baia Vi hai'az: ChiIasco(UTAA-47567-69);
circa 5 km S Chilasco, 1800 m (MVZ 143379); 8 km ESE Chilasco, Finca
Miranda, 6500 ft |198] m| (MVZ 150931); 50,2 km NW El Rancho (UTA
A-=.016); CA-14, 29,0 mi |46,7 km] NW El Rancho (UTA A-.5049); CA-
14, 50,2 km NW El Rancho (UTA A-5015): 4,8 mi (7,7 km] SSE Puruiha,
Plantacion Santa Teresa (UTA A-74 1 7); 9, 1 mi (14,6 km] W Salama (by road
to Puruiha) (UTA A-7432); 2,4 mi ]3,9 km] W Puruiha (UTA A-8502-<')7); 3.5
mi ]5.6 km] W Puruiha (UTA A-30495 larvae); 3.2 km WNW Puruiha (UTA
A-17117-18); 3.5 km W western Puruiha turnoff (UTA A- 17242- 1 7244);
2.7 km W western Puruiha turnoff (UTA A-I7245); 3.8 km W Puruiha,
1536 m (KU 186288-303); 7,7 km SE Puruiha, 1615 m (KU 186304); 3,8
km W Puruiha, 1524 in (KU 190067); 4.2 km W Puruiha, 1524 m (KU
190068-70); 3,4 km W Puruiha, 1524 m (KU 190071); 2,0 km W Puruiha
(UTA A-38145^9); Hwy CA-17 between El Rancho and Coban, km 126
(UTA A-43977-78); 1 km S San Geronimo (UMMZ 84083), El Quich6:
Jovabaj (KU 186305); La Primavera, between Sacapulas and Santa Cruz
Quiche, 6600 ft ]2012 m] (UMMZ 126307), Gi .ufmai a: Amatitlan (UTA
A-.38144); 1 1,2 km SW Guatemala City 4600 ft [1402 m] (KU 97,595-609);
21 km SW Guatemala City, 4480 ft 11366 m] (KU 97610-19]; Guatemala
Citv, /one 10, 4820 ft ]1469 m] (TNHC 31384, 31387, 31390, 31392,
31395, 31399, 31401-02, 31405, 31408, 31416-20, 31422, 31426, 31430-33);
Guatemala City, between zone 5 and zone 15, km 2,5 (UTA A-25824): E
side Guatemala City, zone 16, 1 km N Vista Hermosa III on road to Santa
Rosita (UTA A-25825-32); Santa Catarina Pinula, San Miguel Buena Vista,
1700 m (UTA A-43951, UTA A-47570-74); Guatemala City zone 15, Vista
Hermosa IIL 1510 m (UTA A-28959-60);Parque San Jorge Muxbal 1850 m
(UTA A-32993). HuEHLErtNANt.o: Aguacatan (UMMZ 120046); 2 km NE
Aguacatan, 1640 m (UMMZ 120047-48); 2,8 km E Aguacatan, 1600 m (KU
^8412-13); Huehuetenango, patio of Casa Maryknoll (UMMZ 124382);
22 km SSW Huehuetenango (KU 116959); 3 km W I luehuetenango, 6100
tt ]1859 m] (TNHC 29452-57); at La Libertad, 1700 m (MVZ 143343-57);
San Pedro Necta (UMMZ 130059 larvae); circa 1 km F San Pedro \ecta,
1615 m (UMMZ 119352). Jaiara: Jalapa (TNHC 33666-72); 8.5 km NW
Jalapa (INHC 31442); 7.5 km WSW Jalapa, on road to Miramundo (UTA
A-,391I4 larvae); Jalapa-Miramundo rd, at km 101 (UTA A-,381 18); Falda
Oeste Volcan Jumav (UTA A-47,565); 1,6 mi [2,5 km] NE El Mojon (UTA
A-38127-40); 0,7 nii'l 1 . 1 km] NE EI Mojon (UTA A-38141 ); Aserradero San
Lorenzo (circa 12 air km NNE Jalapa), 1725 m (UMMZ 108000, 106806
] 10 specimens], 106807 ]3 specimens], Proc.reso: Finca Bucaral (UMMZ
10(iS08, 1395 16 larvae), S.u aiii-lqui/: 3 km W Dueiias (TNHC 31378); 1,3
mi ]4,4 km] W Finca San Rafael Urias at Dueiias (TNHC 31344, 31379-80);
San Antonio (CAS 70826-27); Volcan Agua (CAS 70719-825), Honduras:
Kiiiiii. a: water supply area for La Fsperanza (LACM 45247—18) 1,5 mi
NE La Esperanza (LACM 47992-96); I mi NE La Esperanza (LACM
47998); La Esperanza (LACM 47998-48004); 25, 7 km NW La Esperanza,
1340 m (USNM 523689-93); 18,1 km NW La Esperanza, 1740 m (USNM
523694-96); 8,7 km NW La Esperan/a, 1540 m (USNM 523697); Zacate
Blanco, 2020 m (KU 194220-21 ); ca, Miguelito, 60,3 km SF Gracias (Depto.
Lempira), 1310 m (KU 209250, 2092,53), Llmrira: Frandique (LSUMZ
46432, 497.38); above Villa Verde, 1280 m (KU 209240), OtxiTEPEOiiL: 12,5
mi E Nueva Ocot..peque (LACM 47983-85); 6.5 mi E Nueva Ocotepeque
(LACM 47986-91 ); ■•14'29.48'N, 88-46.83'W" (UTA A-52960, ,53662); Belen
Gualcho, 1470 m (KU 194208); El Chaguiton, 1870 m (KU 194209-12;
USNM .523712-13); Fl Volcan, 1760 m (USNM 523714-18),
Bufo /lisiunus: Mexico: Miciioaian: 9 mi on rd between Ri'o
Marque/ <md Cuatro Caminos (KU 62237— 11 ); 1-6 mi S Cuatros Caminos
(LACM 37092-96); 7 mi F: Apal/ing.in, 1 100 fl (UMMZ I1,=;35,5, 112794 ]6
Toads or tiii; Bufo coccifek Complex
27
specimens] ); 1 mi E Apatzingan, 1100 ft (UMMZ 115354 (2 specimens] );
3 mi S Lombardia (UMMZ 121 578): 6.2 mi E Apatzingan, 1100 ft (UMMZ
115353 ]15 specimens], 233723).
Bufo porteri: Honduras: Oim.wacu.a: 10.9 mi N\V Siguatepet|iie
(TNHC 31444); 6.9 mi N\V Siguatepeque, 3820 ft (TNHC 31462, 31466); 7.5
mi NW Siguatepeque, 3500 ft"(KU 97527); 8.8 mi NW Siguatepeque (TNHC
31468); Siguatepeque, 3500 ft (FMNH 4612-13); 9.8 km SW Siguatepeque,
1700 m (USNM 523683); 9.8 km SW Siguatepeque, 1950 m (USNM
523684 );Montana de Comayagua above Ri'o Negro, 1530 m (KU 209247).
Francisco MokazAn: Morizan, 21.4 km SE San Antonio, 4820 ft (TNHC
31446, 31450, 31452, 31454); 17.1 mi S Tegucigalpa, 4900 ft (TNHC 31455);
Cerro Cantagallo, 1840 m (USNM 523686); Monte Crudo, nr EAP, 6000 ft
(AMNH 54757); La Montanuela, above Table Grande, above EAP (AMNH
54758-59): Uyuca, above EAP, 5800 ft (AMNH 54760): Cerro Uyuca, 1900
m (KU 209254): Uvuca, above EAP, above Tatumbia, ca. 5300 ft (AMNH
54761): slopes of Uvuca, 4500-5000 ft (AMNH 54822); 5.5 mi SW Valle
de Angeles (LACM 47970-71): 4.7 mi SW San Juancito (LACM 47972);
8.6 mi NW Comavaguela (LACM 59426-31 ); 5 km W Zambrano, 1635 m
(KU 65373); Cerro Uyuca (LSUMZ 45433, 45439-10, 45456, 46400, 46427,
49737): Cerro La Tigra (LSUMZ 45436, 45444, 45452): Cerro La Tigra, 1840
m (KU 194215-19, 209249); 6 mi NE Escuela Panamericana, Cerro Uyuca,
5400 ft (KU 97514-18); 6 mi NE Escuela Panamericana, Cerro Uyuca,' 5200
ft (KU 97519-26); W slope Cerro Uyuca, 1750 m (KU 103220); W slope
Cerro Uyuca, 1650 m (KU 103221); Parque Nacional La Tigra, above San
Juancito, 2100 m (KU 192294); El Hatillo (LSUMZ 45438, 45441, 45446,
46418; LACM 72072). La P.'\z: Marcala (LSUMZ 46396-98, 46401, 46405,
46407, 46412-14, 46420, 46422-24, 46426, 46428, 46431, 464,34-35, 46442,
46445, 46448-51, 46453-55); Santa Elena, 1750 m (KU 194222); Sierra de
Montecillos, about Tutule, 1750 m (KU 209244).
Biifo signifer: Panama: Canal ZoNt: no further data (TNHC 46261 ).
CuiRiQLii: Cerro Colorado, Escopeta Camp, ca 23 km NNE San Felix, 900 m
(USNM 297511-15, 297517-21); Cerro Bollo, 3.5 km E of Escopeta Camp,
1800-1850 m (USNM 297516, 297522); 7 mi N and 2 mi W David (AMNH
69626); 7 mi E Concepcion (AMNH 69627); Cerro Bollo, 3.5 km E Escopeta
Camp, ca. 1800 m (USNM 297522); 2.5 mi NE David (TNHC 31340-43).
Cocle: Agua Dulce (UMMZ 167438): El Valle, 2000 ft (ANSP 23418-19);
El Valle de Anton (AMNH 59634, 59637); 16 km S and 9 km W Penonome,
30m(KU 1 1 5359-61 ); 3.2 km WAguadulce, 15 m (KU 115362). Herkera:
3 mi SW Pan-Am Hwy on rd past Potuga (UMMZ 167373); Jacinto, 2250
ft (ANSP 22341-14); Vei^guas: 14 km NE Sona, 75 m (KU 95432).
3 2044 072 228 646
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